Aquificales

The ''Aquificota'' phylum is a diverse collection of bacteria that live in harsh environmental settings. The name ''Aquificota'' 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. 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, non-spore-forming rods. They are true bacteria (domain Bacteria) as opposed to the other inhabitants of extreme environments, the Archaea.

Taxonomy

The ''Aquificota'' currently contain 15 genera and 42 validly published species. The phylum comprises a single class and two orders. 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. It is currently classified as a member of Aquificales, but it has shown more physiological similarity to the Desulfobacteriaceae.

Molecular signatures and phylogenetic position

Comparative genomic studies have identified several conserved signature indels (CSIs) that are specific for all species belonging to the phylum ''Aquificota'' and provide potential molecular markers. 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 ''Aquificota'' and Hydrogenothermaceae from all other bacteria. In parallel with the observed CSI distribution, the orders within the ''Aquificota'' 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.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.

Several CSIs have also been identified that are specific for the species from the ''Aquificota'' and provide potential molecular markers for this phylum. Additionally, a 51-amino-acid insertion has been identified in SecA preprotein translocase which is shared by all members of the ''Aquificota'', as well as all members of the order Thermotogales. Phylogenetic studies demonstrated that the presence of the same CSI within these two unrelated groups of bacteria is not due to lateral gene transfer, rather the CSI likely developed independently in these two groups of thermophiles due to selective pressure. The 51 amino acid insertion is located on the surface of SecA near the binding site of ADP/ATP. Molecular dynamic simulations revealed a network water molecules forming an intermediate interaction between residues of the 51 aa CSI and ADP molecules, which serves to stabilize the hydrogen bonds formed between ADP/ATP and the protein. It is suggested that the network of hydrogen bonds formed between the water molecules, CSI residues and ADP/ATP helps to maintain ATP/ADP binding to the SecA protein at high temperatures, which contributes to the bacteria’s overall thermostability.

In the 16S rRNA gene trees, the ''Aquificota'' species branch in the proximity of the phylum Thermotogota (another phylum comprising hyperthermophilic organisms) close to the archaeal-bacterial branch point.Huber, R. and Hannig, M. (2006) Thermotogales. Prokaryotes 7: 899-922. However, a close relationship of the ''Aquificota'' to the Thermotogota and the deep branching of the ''Aquificota'' is not supported by some phylogenetic studies based upon other gene/protein sequences and also by CSIs in several highly conserved universal proteins 16S-23S-5S operons.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. 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, the inference that the ''Aquificota'' 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 Campylobacterota. A specific relationship of the ''Aquificota'' to the Proteobacteria is supported by a two-amino-acid CSI in the protein inorganic pyrophosphatase, which is uniquely found in species from these two phyla. Cavalier-Smith has also suggested that the ''Aquificota'' are closely related to the Proteobacteria. 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. These authors explain the frequently observed grouping of ''Aquificota'' with Campylobacterota as result of frequent horizontal gene transfer due to shared ecological niches.

Along with the Thermotogota, the ''Aquificota'' are thermophilic eubacteria.

Phylogeny

Taxonomy

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

* Class " Thermosulfidibacteria" Cavalier-Smith 2020
** Order " Thermosulfidibacterales" Cavalier-Smith 2020
*** Family " Thermosulfidibacteraceae" Cavalier-Smith 2020
* Class " Aquificia" Oren, Parte & Garrity 2016
** Order Desulfurobacteriales Gupta & Lali 2014
*** Family Desulfurobacteriaceae L'Haridon et al. 2006
** Order Aquificales Reysenbach 2002 em. Gupta & Lali 2013
*** Family Hydrogenothermaceae Eder and Huber 2003
*** Family Aquificaceae Reysenbach 2002

See also

* List of bacterial orders
* List of bacteria genera

References

{{Taxonbar, from=Q597831

Bergey's volume 1
Bacteria phyla