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The Aquificae phylum is a diverse collection of bacteria that live in harsh environmental settings.[1][2] The name 'Aquificae' was given to this phylum based on an early genus identified within this group, Aquifex (“water maker”), which is able to produce water by oxidizing hydrogen.[3] They have been found in springs, pools, and oceans. They are autotrophs, and are the primary carbon fixers in their environments. These bacteria are Gram-negative, nonspore-forming rods.[4] They are true bacteria (domain Bacteria) as opposed to the other inhabitants of extreme environments, the Archaea.

Contents

1 Taxonomy 2 Molecular signatures and phylogenetic position 3 Phylogeny 4 Taxonomy 5 References

Taxonomy[edit] The Aquificae currently contain 15 genera and 42 validly published species.[5] The phylum comprises a single class and two orders.[6][7] Aquificales consists of the families Aquificaceae and Hydrogenothermaceae, while the Desulfurobacteriaceae are the only family within the Desulfobacteriales. Thermosulfidibacter takaii is not assigned to a family within the phylum based on its phylogenetic distinctness from both orders.[8] It is currently classified as a member of Aquificales, but it has shown more physiological similarity to the Desulfobacteriaceae. Molecular signatures and phylogenetic position[edit] Comparative genomic studies have identified several conserved signature indels (CSIs) that are specific for all species belonging to the phylum Aquificae and provide potential molecular markers.[7] The order Aquificales can be distinguished from Desulfobacteriales by several CSIs across different proteins that are specific for each group. Additional CSIs have been found at the family level, and can be used to demarcate Aquificae and Hydrogenothermaceae from all other bacteria.[7] In parallel with the observed CSI distribution, the orders within the Aquificae are also physiologically distinct from one another. Members of the Desulfurobacteriales are strict anaerobes that exclusively oxidize hydrogen for energy, whereas those belonging to the Aquificales are microaerophilic, and capable of oxidizing other compounds (such as sulfur or thiosulfate) in addition to hydrogen.[9][10][11] Several CSIs have also been identified that are specific for the species from the Aquificae and provide potential molecular markers for this phylum.[1] Additionally, a 51-amino-acid insertion has been identified in SecA preprotein translocase which is shared by various members of the Aquificae, as well as two Thermotoga
Thermotoga
species. In the 16S rRNA gene trees, the Aquificae species branch in the proximity of the phylum Thermotogae (another phylum comprising hyperthermophilic organisms) close to the archaeal-bacterial branch point.[12][10] However, a close relationship of the Aquificae to the Thermotogae and the deep branching of the Aquificae is not supported by some phylogenetic studies based upon other gene/protein sequences[13][14][15][16] and also by CSIs in several highly conserved universal proteins.[17] Thus, the presence of the insertion in the Thermotoga
Thermotoga
species may be due to a horizontal gene transfer.[1] The deep branching of Aquificae species in the rRNA gene tree was ascribed to be an artefact resulting from the very high G+C content of their 16S-23S-5S operons.[17] In contrast to the very high G+C content of their rRNAs (i.e. more than 62%), which is required for stability of their secondary structures at high growth temperatures,[18] the inference that the Aquificae do not constitute a deep-branch lineage is also independently strongly supported by CSIs in a number of important proteins (viz. Hsp70, Hsp60, RpoB, RpoB and AlaRS), which support its placement in the proximity of the phylum Proteobacteria, particularly the Epsilonproteobacteria.[17] A specific relationship of the Aquificae to the Proteobacteria
Proteobacteria
is supported by a two-amino-acid CSI in the protein inorganic pyrophosphatase, which is uniquely found in species from these two phyla.[17] Cavalier-Smith
Cavalier-Smith
has also suggested that the Aquificae are closely related to the Proteobacteria.[19] In contrast to the above cited analyses that are based on a few indels or on single genes, analyses on informational genes, which appeared to be less often transferred to the Aquifex lineage than noninformational genes, most often placed the Aquificales close to the Thermotogales.[20] These authors explain the frequently observed grouping of Aquificae with Epsilonproteobacteria
Epsilonproteobacteria
as result of frequent horizontal gene transfer due to shared ecological niches. Along with the Thermotogae, the Aquificae are thermophilic eubacteria.[2] Phylogeny[edit] The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN) [5] and National Center for Biotechnology Information (NCBI)[21] and the phylogeny is based on 16S rRNA-based LTP release 123 by The All-Species Living Tree Project
The All-Species Living Tree Project
[22] [23]

Thermosulfidibacter takaii

Desulfurobacteriaceae

Phorcysia thermohydrogeniphila

Desulfurobacterium thermolithotrophum

Desulfurobacterium atlanticum

Desulfurobacterium pacificum

Thermovibrio

T. ruber

Balnearium lithotrophicum

T. ammonificans

T. guaymasensis

Aquificales

Hydrogenothermaceae

Persephonella hydrogeniphila

Persephonella marina

Persephonella guaymasensis

Venenivibrio
Venenivibrio
stagnispumantis

Sulfurihydrogenibium

S. yellowstonense

S. kristjanssonii

S. subterraneum

S. azorense

S. rodmanii

Aquificaceae

Hydrogenobaculum acidophilum

Hydrogenobacter

H. subterraneus

H. hydrogenophilus

H. thermophilus

Thermocrinis minervae

Thermocrinis ruber

Thermocrinis albus

Aquifex pyrophilus

Hydrogenivirga

H. calditorris

H. okinawensis

Note: ♠ Strain found at the National Center for Biotechnology Information (NCBI) but not listed in the List of Prokaryotic names with Standing in Nomenclature (LPSN) Taxonomy[edit]

Genus Thermosulfidibacter Nunoura et al. 2008

Species Thermosulfidibacter takaii Nunoura et al. 2008

Order Desulfurobacteriales Gupta & Lali 2014

Family Desulfurobacteriaceae L'Haridon et al. 2006

Genus Balnearium Takai et al. 2003

Species Balnearium lithotrophicum Takai et al. 2003

Genus Desulfurobacterium L'Haridon et al. 1998 emend. L'Haridon et al. 2006

Species "D. crinifex" ♠ Alain et al. 2003 Species D. atlanticum L'Haridon et al. 2006 Species D. pacificum L'Haridon et al. 2006 Species D. thermolithotrophum L'Haridon et al. 1998 (type sp.)

Genus Phorcysia Pérez-Rodríguez et al. 2012

Species Phorcysia thermohydrogeniphila Pérez-Rodríguez et al. 2012

Genus Thermovibrio Huber et al. 2002

Species T. ammonificans Vetriani et al. 2004 Species T. guaymasensis L'Haridon et al. 2006 Species T. ruber Huber et al. 2002 (type sp.)

Order Aquificales Reysenbach 2002 emend. Gupta & Lali 2014

Family Aquificaceae Reysenbach 2002

Genus Aquifex Huber and Stetter 1992

Species A. pyrophilus Huber and Stetter 1992 (type sp.) Species "A. aeolicus" ♠ Huber and Stetter 2001

Genus Hydrogenivirga Nakagawa et al. 2004

Species H. calditorris Nakagawa et al. 2004 (type sp.) Species H. okinawensis Nunoura et al. 2008

Genus Hydrogenobacter Kawasumi et al. 1984

Species H. hydrogenophilus (Kryukov et al. 1984) Stöhr et al. 2001 Species H. subterraneus Takai et al. 2001 Species H. thermophilus Kawasumi et al. 1984 (type sp.)

Genus Hydrogenobaculum Stöhr et al. 2001

Species Hydrogenobaculum acidophilum (Shima and Suzuki 1993) Stöhr et al. 2001

Genus Thermocrinis Huber et al. 1999

Species T. albus Eder and Huber 2002 Species T. jamiesonii Dodsworth et al. 2015 Species T. minervae Huber et al. 1999 Species T. ruber Huber et al. 1999 (type sp.)

Family Hydrogenothermaceae Eder and Huber 2003

Genus Hydrogenothermus Stöhr et al. 2001

Species Hydrogenothermus marinus Stöhr et al. 2001

Genus Persephonella Götz et al. 2002

Species P. hydrogeniphila Nakagawa et al. 2003 Species P. marina Götz et al. 2002 (type sp.) Species P. guaymasensis Götz et al. 2002

Genus Sulfurihydrogenibium Takai et al. 2003 emend. O'Neill et al. 2008

Species S. azorense Aguiar et al. 2004 emend. Nakagawa et al. 2005 Species S. kristjanssonii Flores et al. 2008 Species S. rodmanii O'Neill et al. 2008 Species S. subterraneum Takai et al. 2003 emend. Nakagawa et al. 2005 (type sp.) Species S. yellowstonense Nakagawa et al. 2005

Genus Venenivibrio
Venenivibrio
Hetzer et al. 2008

Species Venenivibrio stagnispumantis
Venenivibrio stagnispumantis
Hetzer et al. 2008

References[edit]

^ a b c Griffiths E, Gupta RS (January 2006). "Molecular signatures in protein sequences that are characteristics of the phylum Aquificae". Int. J. Syst. Evol. Microbiol. 56 (Pt 1): 99–107. doi:10.1099/ijs.0.63927-0. PMID 16403873.  ^ a b Horiike T, Miyata D, Hamada K, et al. (January 2009). "Phylogenetic construction of 17 bacterial phyla by new method and carefully selected orthologs". Gene. 429 (1–2): 59–64. doi:10.1016/j.gene.2008.10.006. PMC 2648810 . PMID 19000750.  ^ Huber R, Wilharm T, Huber D, Trincone A, Burggaf S, Konig H, Reinhard R, Rockinger I, Fricke H, Stetter K (1992). "Aquifex pyrophilus gen. nov. sp. nov., Represents a Novel Group of Marine Hyperthermophilic Hydrogen-Oxidizing Bacteria". Syst. Appl. Microbiol. 15: 340–351. doi:10.1016/S0723-2020(11)80206-7.  ^ L'Haridon, Reysenbach AL, Tindall BJ, Schönheit P, Banta A, Johnsen U, Schumann P, Gambacorta A, Stackebrandt E, Jeanthon C (December 2006). " Desulfurobacterium atlanticum sp. nov., Desulfurobacterium pacificum sp. nov. and Thermovibrio guaymasensis sp. nov., three thermophilic members of the Desulfurobacteriaceae fam. nov., a deep branching lineage within the bacteria". Int. J. Syst. Evol. Microbiol. 56 (Pt 12): 2843–2852. doi:10.1099/ijs.0.63994-0. PMID 17158986.  ^ a b J.P. Euzéby. "Aquificae". List of Prokaryotic names with Standing in Nomenclature (LPSN). Archived from the original on 2011-06-13. Retrieved 2016-09-09.  ^ Oren A, Garrity GM (2015). "List of new names and new combinations previously effectively, but not validly, published" (PDF). Int. J. Syst. Evol. Microbiol. 65 (7): 2017–2025. doi:10.1099/ijs.0.000317.  ^ a b c Gupta RS, Lali R (September 2013). "Molecular signatures for the phylum Aquificae and its different clades: proposal for division of the phylum Aquificae into the emended order Aquificales, containing the families Aquificaceae and Hydrogenothermaceae, and a new order Desulfurobacteriales ord. nov., containing the family Desulfurobacteriaceae". Antonie van Leeuwenhoek. 104 (3): 349–368. doi:10.1007/s10482-013-9957-6. PMID 23812969.  ^ Nunoura T, Oida H, Miyazaki M, Suzuki Y (March 2008). "Thermosulfidibacter takaii gen. nov., sp. nov., a thermophilic, hydrogen-oxidizing, sulfur-reducing chemolithoautotroph isolated from a deep-sea hydrothermal field in the Southern Okinawa Trough". Int. J. Syst. Evol. Microbiol. 58 (Pt 3): 659–665. doi:10.1099/ijs.0.65349-0. PMID 18319474.  ^ Guiral M, Prunetti L, Aussignargues C, Ciaccafava A, Infossi P, Ilbert M, Lojou E, Giudici-Orticoni MT (2012). "The hyperthermophilic bacterium Aquifex aeolicus: from respiratory pathways to extremely resistant enzymes and biotechnological applications". Adv. Microb. Physiol. 61: 125–194. doi:10.1016/B978-0-12-394423-8.00004-4. PMID 23046953.  ^ a b Reysenbach, A.-L. (2001) Phylum BII. Thermotogae phy. nov. In: Bergey's Manual of Systematic Bacteriology, pp. 369-387. Eds D. R. Boone, R. W. Castenholz. Springer-Verlag: Berlin. ^ Gupta, RS (2014) The Phylum Aquificae. The Prokaryotes 417-445. Springer Berlin Heidelberg. ^ Huber, R. and Hannig, M. (2006) Thermotogales. Prokaryotes 7: 899-922. ^ Klenk, H. P., Meier, T. D., Durovic, P. and others (1999) RNA polymerase of Aquifex pyrophilus: Implications for the evolution of the bacterial rpoBC operon and extremely thermophilic bacteria. J Mol Evol 48: 528-541. ^ Gupta, R. S. (2000) The phylogeny of Proteobacteria: relationships to other eubacterial phyla and eukaryotes. FEMS Microbiol Rev 24: 367-402. ^ Ciccarelli, F. D., Doerks, T., von Mering, C., Creevey, C. J., Snel, B., and Bork, P. (2006) Toward automatic reconstruction of a highly resolved tree of life. Science 311: 1283-1287. ^ Di Giulio, M. (2003) The universal ancestor was a thermophile or a hyperthermophile: Tests and further evidence. J Theor Biol 221: 425-436. ^ a b c d Griffiths, E. and Gupta, R. S. (2004) Signature sequences in diverse proteins provide evidence for the late divergence of the order Aquificales. International Microbiol 7: 41-52. ^ Meyer, T. E. and Bansal, A. K. (2005) Stabilization against hyperthermal denaturation through increased CG content can explain the discrepancy between whole genome and 16S rRNA analyses. Biochemistry 44: 11458-11465. ^ Catalogue of Organisms: Standing the Heat ^ Boussau B, Guéguen L, Gouy M. Accounting for horizontal gene transfers explains conflicting hypotheses regarding the position of Aquificales in the phylogeny of Bacteria. BMC Evol Biol. 2008 Oct 3;8:272. doi:10.1186/1471-2148-8-272. ^ Sayers; et al. "Aquificae". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 2016-09-09.  ^ All-Species Living Tree Project."16S rRNA-based LTP release 123 (full tree)" (PDF). Silva Comprehensive Ribosomal RNA Database. Retrieved 2016-09-09.  ^ Ludwig, W.; Euzéby, J. & Whitman W.B. (2008). "Bergey's Taxonomic Outlines: Volume 4 - Draft Taxonomic Outline of the Bacteroidetes, Planctomycetes, Chlamydiae, Spirochaetes, Fibrobacteres, Fusobacteria, Acidobacteria, Verrucomicrobia, Dictyoglomi, and Gemmatimonadetes" (PDF). Bergey's Manual Trust: 15. 

v t e

Prokaryotes: Bacteria
Bacteria
classification (phyla and orders)

Domain Archaea Bacteria Eukaryota (Supergroup Plant Hacrobia Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta

Animal Fungi)

G-/ OM

Terra-/ Glidobacteria (BV1)

Eobacteria

Deinococcus–Thermus

Deinococcales Thermales

Chloroflexi

Anaerolineales Caldilineales Chloroflexales Herpetosiphonales Dehalococcoidales Ktedonobacterales Thermogemmatisporales Thermomicrobiales Sphaerobacterales

other glidobacteria

Thermodesulfobacteria thermophiles

Aquificae Thermotogae

Cyanobacteria

Proteobacteria (BV2)

Alpha

Caulobacterales Kiloniellales Kordiimonadales Magnetococcales Parvularculales Rhizobiales Rhodobacterales Rhodospirillales Rickettsiales Sneathiellales Sphingomonadales

Beta

Burkholderiales Hydrogenophilales Methylophilales Neisseriales Nitrosomonadales Procabacteriales Rhodocyclales

Gamma

Acidithiobacillales Aeromonadales Alteromonadales Cardiobacteriales Chromatiales Enterobacteriales Legionellales Methylococcales Oceanospirillales Orbales Pasteurellales Pseudomonadales Salinisphaerales Thiotrichales Vibrionales Xanthomonadales

Delta

Bdellovibrionales Desulfarculales Desulfobacterales Desulfovibrionales Desulfurellales Desulfuromonadales Myxococcales Syntrophobacterales Syntrophorhabdales

Epsilon

Campylobacterales Nautiliales

Zeta

Mariprofundales

BV4

Spirochaetes

Spirochaetes

Sphingobacteria (FCB group)

Fibrobacteres Chlorobi

Chlorobiales Ignavibacteriales

Bacteroidetes

Bacteroidales Cytophagales Flavobacteriales Sphingobacteriales

Planctobacteria/ (PVC group)

Chlamydiae Lentisphaerae

Lentisphaerales Oligosphaerales Victivallales

Planctomycetes

Phycisphaerales Planctomycetales

Verrucomicrobia

Puniceicoccales Opitutales Chthoniobacterales Verrucomicrobiales

"Poribacteria"

Other GN

Acidobacteria

Acidobacteriales Acanthopleuribacterales Holophagales Solibacterales

Armatimonadetes

Armatimonadales Chthonomonadales Fimbriimonadales

Caldiserica Chrysiogenetes Deferribacteres Dictyoglomi Elusimicrobia Fusobacteria Gemmatimonadetes Nitrospirae Synergistetes

G+/ no OM

Firmicutes (BV3)

Bacilli

Bacillales Lactobacillales

Clostridia

Clostridiales Halanaerobiales Thermoanaerobacteriales Natranaerobiales

Erysipelotrichia

Erysipelotrichiales

Thermolithobacteria

Thermolithobacterales

Tenericutes/ Mollicutes

Mycoplasmatales Entomoplasmatales Anaeroplasmatales Acholeplasmatales Haloplasmatales

Negativicutes

Selenomonadales

Actinobacteria (BV5)

Actinobacteria

Actinomycetales Bifidobacteriales

Acidimicrobiia

Acidimicrobiales

Coriobacteriidae

Coriobacteriales

Nitriliruptoria

Euzebyales Nitriliruptorales

Rubrobacteria

Gaiellales Rubrobacterales Thermoleophilales Solirubrobacterales

Incertae sedis

†Archaeosphaeroides †Eobacterium †Leptotrichites

Source: Bergey's Manual (2001–2012). Alternative views: Wikispecies.

Taxon identifiers

Wd: Q597831 EoL: 7855 ITIS: 956100

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