Pelagibacterales

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"Pelagibacterales"
Deposition velocity versus cell diameters-3.jpg
Microscopy image of SAR11
Scientific classification
Domain:
Phylum:
Class:
Subclass:
Order:
"Pelagibacterales"

Grote et al. 2012 [1]
Families
Synonyms [1]
  • SAR11 Clade

The Pelagibacterales are an order in the Alphaproteobacteria composed of free-living marine bacteria that make up roughly one in three cells at the ocean's surface. [2] [3] [4] Overall, members of the Pelagibacterales are estimated to make up between a quarter and a half of all prokaryotic cells in the ocean. [5]

Contents

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. [4] It includes the highly abundant marine species Pelagibacter ubique .

Bacteria in this order are unusually small. [6] Due to their small genome size and limited metabolic function, Pelagibacterales have become a model organism for 'streamlining theory'. [5]

P. ubique and related species are oligotrophs (scavengers) and feed on dissolved organic carbon and nitrogen. [3] They are unable to fix carbon or nitrogen, but can perform the TCA cycle with glyoxylate bypass and are able to synthesise all amino acids except glycine, [7] as well as some cofactors. [8] They also have an unusual and unexpected requirement for reduced sulfur. [9]

P. ubique and members of the oceanic subgroup I possess gluconeogenesis, but not a typical glycolysis pathway, whereas other subgroups are capable of typical glycolysis. [10]

Unlike Acaryochloris marina , P. ubique is not photosynthetic — specifically, it does not use light to increase the bond energy of an electron pair — but it does possess proteorhodopsin (including retinol biosynthesis) for ATP production from light. [11]

SAR11 bacteria are responsible for much of the dissolved methane in the ocean surface. They extract phosphate from methylphosphonic acid. [12]

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. [13]

Subgroups

Currently, the order is divided into five subgroups: [14]

The above results in a cladogram of the Pelagibacterales as follows:

Subgroup Ia (named  Pelagibacteraceae, includes  Pelagibacter )

Subgroup Ib

Subgroup II

Subgroup IIIa

Subgroup IIIb

Subgroup IV (named LD12 clade, includes SAR11 bacteria)

Subgroup V (includes α-proteobacterium HIMB59)

Phylogenetic placement and endosymbiotic theory

A 2011 study by researchers of the University of Hawaiʻi at Mānoa and Oregon State University, indicated that SAR11 could be the ancestor of mitochondria in most eukaryotic cells. [2] However, this result could represent a tree reconstruction artifact due to compositional bias. [16]

Schematic ribosomal RNA phylogeny of Alphaproteobacteria
   Magnetococcidae   

  Magnetococcus marinus

   Caulobacteridae   

  Rhodospirillales, Sphingomonadales,
  Rhodobacteraceae, Hyphomicrobiales, etc.

  Holosporales

   Rickettsidae   
   Pelagibacterales   
   Pelagibacteraceae   

  Pelagibacter

  Subgroups Ib, II, IIIa, IIIb, IV and V

   Rickettsiales   

  Proto-mitochondria

   Anaplasmataceae   

  Ehrlichia

  Anaplasma

  Wolbachia

  Neorickettsia

   Midichloriaceae   

  Midichloria

   Rickettsiaceae   

  Rickettsia

  Orientia

The cladogram of Rickettsidae has been inferred by Ferla et al. [4] from the comparison of 16S + 23S ribosomal RNA sequences.

Related Research Articles

<i>Candidatus Pelagibacter communis</i> Species of bacterium

"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 clade in the phylum Alphaproteobacteria. SAR11 members are highly dominant organisms found in both salt and fresh water worldwide and were originally known only from their rRNA genes, first identified in the Sargasso Sea in 1990 by Stephen Giovannoni's laboratory at Oregon State University and later found in oceans worldwide. "Ca. P. communis" and its relatives may be the most abundant organisms in the ocean, and quite possibly the most abundant bacteria in the entire world. It can make up about 25% of all microbial plankton cells, and in the summer they may account for approximately half the cells present in temperate ocean surface water. The total abundance of "Ca. P. communis" and relatives is estimated to be about 2 × 1028 microbes.

<span class="mw-page-title-main">Rickettsiales</span> Order of bacteria

The Rickettsiales, informally called rickettsias, are an order of small Alphaproteobacteria. They are obligate intracellular parasites, and some are notable pathogens, including Rickettsia, which causes a variety of diseases in humans, and Ehrlichia, which causes diseases in livestock. Another genus of well-known Rickettsiales is the Wolbachia, which infect about two-thirds of all arthropods and nearly all filarial nematodes. Genetic studies support the endosymbiotic theory according to which mitochondria and related organelles developed from members of this group.

<span class="mw-page-title-main">Proteorhodopsin</span> Family of transmembrane proteins

Proteorhodopsin 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 eukaryotes (protae), but was first discovered in bacteria.

An exopolymer is a biopolymer that is secreted by an organism into the environment. These exopolymers include the biofilms produced by bacteria to anchor them and protect them from environmental conditions. One type of expolymer, Transparent Exopolymers (TEP), found in both marine and aquatic ecosystems, are planktonic acidic polysaccharides of a gel-like consistency, originally defined by their ability to be stained visible by acidic Alcian Blue. Their free-floating characteristic sets TEPs aside from other extracellular polymeric substance subgroups where exopolymers exists as cell coating, dissolved slime or as part of biofilm matrices.

Photoheterotrophs are heterotrophic phototrophs—that is, they are organisms that use light for energy, but cannot use carbon dioxide as their sole carbon source. Consequently, they use organic compounds from the environment to satisfy their carbon requirements; these compounds include carbohydrates, fatty acids, and alcohols. Examples of photoheterotrophic organisms include purple non-sulfur bacteria, green non-sulfur bacteria, and heliobacteria. These microorganisms are ubiquitous in aquatic habitats, occupy unique niche-spaces, and contribute to global biogeochemical cycling. Recent research has also indicated that the oriental hornet and some aphids may be able to use light to supplement their energy supply.

<span class="mw-page-title-main">Alphaproteobacteria</span> Class of bacteria

Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota. The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse and possess few commonalities, but nevertheless share a common ancestor. Like all Proteobacteria, its members are gram-negative and some of its intracellular parasitic members lack peptidoglycan and are consequently gram variable.

"Candidatus Midichloria" is a candidatus genus of Gram-negative, non-endospore-forming bacteria, with a bacillus shape around 0.45 µm in diameter and 1.2 µm in length. First described in 2004 with the temporary name IricES1, "Candidatus Midichloria" species are symbionts of several species of hard ticks. They live in the cells of the ovary of the females of this tick species. These bacteria have been observed in the mitochondria of the host cells, a trait that has never been described in any other symbiont of animals.

<span class="mw-page-title-main">Gammaproteobacteria</span> Class of bacteria

Gammaproteobacteria is a class of bacteria in the phylum Pseudomonadota. It contains about 250 genera, which makes it the most genus-rich taxon of the Prokaryotes. Several medically, ecologically, and scientifically important groups of bacteria belong to this class. It is composed by all Gram-negative microbes and is the most phylogenetically and physiologically diverse class of Proteobacteria.

Mycoplasma laboratorium or Synthia refers to a synthetic strain of bacterium. The project to build the new bacterium has evolved since its inception. Initially the goal was to identify a minimal set of genes that are required to sustain life from the genome of Mycoplasma genitalium, and rebuild these genes synthetically to create a "new" organism. Mycoplasma genitalium was originally chosen as the basis for this project because at the time it had the smallest number of genes of all organisms analyzed. Later, the focus switched to Mycoplasma mycoides and took a more trial-and-error approach.

<span class="mw-page-title-main">Bacterioplankton</span> Bacterial component of the plankton that drifts in the water column

Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλανκτος, meaning "wanderer" or "drifter", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and freshwater.

<span class="mw-page-title-main">IMES-2 RNA motif</span>

The IMES-2 RNA motif is a conserved RNA structure that was identified by a study based on metagenomics and bioinformatics, and the underlying RNA sequences were identified independently by a similar earlier study. These RNAs are present in environmental sequences, and when discovered were not known to be present in any cultivated species. However, an IMES-2 RNA has been detected in alphaproteobacterium HIMB114, which is classified in the SAR11 clade of marine bacteria. This finding fits with earlier predictions that species that use IMES-2 RNAs are most closely related to alphaproteobacteria. IMES-2 RNAs are exceptionally abundant, as twice as many IMES-2 RNAs were found as ribosomes in RNAs sampled from the Pacific Ocean. Only two bacterial RNAs are known to be more highly transcribed than ribosomes.

<span class="mw-page-title-main">Bacterial phyla</span> Phyla or divisions of the domain Bacteria

Bacterial phyla constitute the major lineages of the domain Bacteria. While the exact definition of a bacterial phylum is debated, a popular definition is that a bacterial phylum is a monophyletic lineage of bacteria whose 16S rRNA genes share a pairwise sequence identity of ~75% or less with those of the members of other bacterial phyla.

HTVC010P is a virus which was discovered by Stephen Giovannoni and colleagues at Oregon State University. The Economist reports that a February 2013 paper in Nature says that "it probably really is the commonest organism on the planet". It is a bacteriophage that infects the extremely abundant bacteria Pelagibacter ubique in the Pelagibacterales order.

Lentisphera araneosa is a marine bacteria strain in the bacterial phylum Lentisphaerota. They are able to produce viscous transparent exopolymers and grow attached to each other by the polymer in a three-dimensional configuration. They are part of the natural surface bacterial population in the Atlantic and Pacific oceans. They are less than 1% of the total bacterial community. This species is gram negative, non-motile, non-pigmented, aerobic, chemoheterotrophic, and facultatively oligotrophic sphere-shaped. Its genome has been sequenced.

The Pelagibacteraceae are a family in the Alphaproteobacteria composed of free-living marine bacteria.

<span class="mw-page-title-main">Kill the Winner hypothesis</span> Microbiological population model hypothesis

The "Kill the Winner" hypothesis (KtW) is an ecological model of population growth involving prokaryotes, viruses and protozoans that links trophic interactions to biogeochemistry. The model is related to the Lotka–Volterra equations. It assumes that prokaryotes adopt one of two strategies when competing for limited resources: priority is either given to population growth ("winners") or survival ("defenders"). As "winners" become more abundant and active in their environment, their contact with host-specific viruses increases, making them more susceptible to viral infection and lysis. Thus, viruses moderate the population size of "winners" and allow multiple species to coexist. Current understanding of KtW primarily stems from studies of lytic viruses and their host populations.

Bacterioplankton counting is the estimation of the abundance of bacterioplankton in a specific body of water, which is useful information to marine microbiologists. Various counting methodologies have been developed over the years to determine the number present in the water being observed. Methods used for counting bacterioplankton include epifluorescence microscopy, flow cytometry, measures of productivity through frequency of dividing cells (FDC), thymidine incorporation, and leucine incorporation.

Genomic streamlining is a theory in evolutionary biology and microbial ecology that suggests that there is a reproductive benefit to prokaryotes having a smaller genome size with less non-coding DNA and fewer non-essential genes. There is a lot of variation in prokaryotic genome size, with the smallest free-living cell's genome being roughly ten times smaller than the largest prokaryote. Two of the bacterial taxa with the smallest genomes are Prochlorococcus and Pelagibacter ubique, both highly abundant marine bacteria commonly found in oligotrophic regions. Similar reduced genomes have been found in uncultured marine bacteria, suggesting that genomic streamlining is a common feature of bacterioplankton. This theory is typically used with reference to free-living organisms in oligotrophic environments.

Rappephyceae, or Rappemonads, are a small family of protists first described in 2011, of uncertain phylogenic affinity. It has been discussed as a possible member of a larger clade Haptophyta. This newly identified taxonomic class of phytoplankton are named after a professor from the Hawai’i institute of marine biology, known as Michael Rappé. Rappé discovered these phytoplankton in the Atlantic Ocean and published his findings on their DNA in 1998. Current research has shown that these organisms provide an immense amount of nutritional organic molecules, such as oxygen, for other organisms using biochemical processes like photosynthesis and carbon fixation.

<span class="mw-page-title-main">Marine prokaryotes</span> Marine bacteria and marine archaea

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 life forms can be divided into prokaryotes and eukaryotes. Eukaryotes are organisms whose cells have a nucleus enclosed within membranes, whereas prokaryotes are the organisms that do not have a nucleus enclosed within a membrane. The three-domain system of classifying life adds another division: the prokaryotes are divided into two domains of life, the microscopic bacteria and the microscopic archaea, while everything else, the eukaryotes, become the third domain.

References

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