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The Pelagibacterales are an order in the
Alphaproteobacteria Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota (formerly Proteobacteria). The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse and p ...
composed of free-living
marine bacteria Marine prokaryotes are marine bacteria and marine archaea. They are defined by their habitat as prokaryotes that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. All cellular ...
that make up roughly one in three cells at the ocean's surface. Overall, members of the ''Pelagibacterales'' are estimated to make up between a quarter and a half of all
prokaryotic A prokaryote () is a Unicellular organism, single-celled organism that lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek language, Greek wikt:πρό#Ancient Greek, πρό (, 'before') a ...
cells in the ocean. Initially, this taxon was known solely by metagenomic data and was known as the SAR11 clade. It was first placed in the Rickettsiales, but was later raised to the rank of order, and then placed as sister order to the Rickettsiales in the subclass Rickettsidae. It includes the highly abundant marine species ''
Pelagibacter ubique "''Candidatus'' Pelagibacter", with the single species "''Ca.'' P. communis", was isolated in 2002 and given a specific name, although it has not yet been described as required by the bacteriological code. It is an abundant member of the SAR11 c ...
''. Bacteria in this order are unusually small. Due to their small genome size and limited metabolic function, ''Pelagibacterales'' have become a model organism for ' streamlining theory'. ''P. ubique'' and related species are
oligotroph An oligotroph is an organism that can live in an environment that offers very low levels of nutrients. They may be contrasted with copiotrophs, which prefer nutritionally rich environments. Oligotrophs are characterized by slow growth, low rates of ...
s (scavengers) and feed on dissolved organic carbon and nitrogen. They are unable to fix carbon or nitrogen, but can perform the
TCA cycle The citric acid cycle (CAC)—also known as the Krebs cycle or the TCA cycle (tricarboxylic acid cycle)—is a series of chemical reactions to release stored energy through the oxidation of acetyl-CoA derived from carbohydrates, fats, and protein ...
with glyoxylate bypass and are able to synthesise all amino acids except glycine, as well as some cofactors. They also have an unusual and unexpected requirement for reduced sulfur. ''P. ubique'' and members of the oceanic subgroup I possess
gluconeogenesis Gluconeogenesis (GNG) is a metabolic pathway that results in the generation of glucose from certain non-carbohydrate carbon substrates. It is a ubiquitous process, present in plants, animals, fungi, bacteria, and other microorganisms. In vertebrat ...
, but not a typical
glycolysis Glycolysis is the metabolic pathway that converts glucose () into pyruvate (). The free energy released in this process is used to form the high-energy molecules adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide (NADH ...
pathway, whereas other subgroups are capable of typical glycolysis. Unlike '' Acaryochloris marina'', ''P. ubique'' is not
photosynthetic Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...
— specifically, it does not use light to increase the bond energy of an electron pair — but it does possess
proteorhodopsin Proteorhodopsin (also known as pRhodopsin) is a family of transmembrane proteins that use retinal as a chromophore for light-mediated functionality, in this case, a proton pump. pRhodopsin is found in marine planktonic bacteria, archaea and e ...
(including
retinol Retinol, also called vitamin A1, is a fat-soluble vitamin in the vitamin A family found in food and used as a dietary supplement. As a supplement it is used to treat and prevent vitamin A deficiency, especially that which results in xerophtha ...
biosynthesis) for ATP production from light. SAR11 bacteria are responsible for much of the dissolved
methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane on Eart ...
in the ocean surface. They extract phosphate from
methylphosphonic acid Methylphosphonic acid is an organophosphorus compound with the chemical formula CH3P(O)(OH)2. The phosphorus center is tetrahedral and is bonded to a methyl group, two OH groups and an oxygen. Methylphosphonic acid is a white, non-volatile solid t ...
. Although the taxon derives its name from the type species ''P. ubique'' (status'' Candidatus'' species), this species has not yet been validly published, and therefore neither the order name nor the species name has official taxonomic standing.


Subgroups

Currently, the order is divided into five subgroups: * Subgroup Ia, open ocean, crown group — includes ''P. ubique'' HTCC1062 * Subg * Subgroup II, coastal, basal to Ia + Ib * Subgroup III, brackish, basal to I + II along with its sister clade IV * Subgroup IV, also known as the LD12 clade, freshwater * Subgroup V, which includes alphaproteobacterium HIMB59, basal to the remainder The above results in a cladogram of the Pelagibacterales as follows:


Phylogenetic placement and endosymbiotic theory

A 2011 study by researchers of the
University of Hawaiʻi at Mānoa The University of Hawaii at Mānoa (University of Hawaii—Mānoa, UH Mānoa, Hawai'i, or simply UH) is a Public university, public Land-grant university, land-grant research university in Manoa, Mānoa, a neighborhood in Honolulu, Hawaii. It ...
and
Oregon State University Oregon State University (OSU) is a public land-grant, research university in Corvallis, Oregon. OSU offers more than 200 undergraduate-degree programs along with a variety of graduate and doctoral degrees. It has the 10th largest engineering co ...
, indicated that SAR11 could be the ancestor of
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
in most eukaryotic cells. However, this result could represent a tree reconstruction artifact due to compositional bias.


References

{{Taxonbar, from=Q7388740 Alphaproteobacteria