Endozoicomonas
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''Endozoicomonas'' is a genus of
Gram-negative Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wa ...
,
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cel ...
or
facultatively anaerobic A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...
,
chemoorganotrophic Primary nutritional groups are groups of organisms, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the ...
,
rod-shaped A bacillus (), also called a bacilliform bacterium or often just a rod (when the context makes the sense clear), is a rod-shaped bacterium or archaeon. Bacilli are found in many different taxonomic groups of bacteria. However, the name '' Baci ...
,
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 ...
from the family of
Hahellaceae The ''Hahellaceae'' are a family of Pseudomonadota in the order of Oceanospirillales The Oceanospirillales are an order of Pseudomonadota with ten families. Description Bacteria in the ''Oceanospirillales'' are metabolically and morphologic ...
. ''Endozoicomonas'' are symbionts of marine animals.


Scientific History

The genus was firstly proposed in 2007 after isolating an unknown Gammaproteobacteria from the sea slug ''
Elysia ornata ''Elysia ornata'', commonly known as ornate elysia or ornate leaf slug, is a species of sea slug, a marine (ocean), marine gastropod mollusk. This sea slug superficially resembles a nudibranch, yet it does not belong to that suborder of gastropo ...
''. Called '' E. numazuensis,'' it was the first of many diverse species now known, and was collected from seawater off the coast of Izu-Miyake Island, Japan, at a depth of 15 m. Many new species have been identified after this: * In 2010, researchers from different Asian universities isolated '' E. montiporae'' from the encrustating pores of coral species ''
Montipora aequituberculata ''Montipora aequituberculata'' is a species of stony coral in the family Acroporidae. It is a common coral in shallow water in the Indo-Pacific region. Distribution and habitat ''Montipora aequituberculata'' is native to the Indo-Pacific region ...
,'' in Taiwan, and a scientific team of the University of Queensland, Australia, discovered many unidentified Gammaproteobacteria symbionts were closely related to ''Endozoicomonas'' as well during that same year. * More or less in the same period '' E. arenosclerae'' was isolated from the endemic marine sponge of Rio de Janeiro '' Arenosclera brasiliensis'' and, both '' E.eunicecola'' and '' E.gorgoniicola'' were isolated from the octocorals '' Eunicea fusca'' and ''
Plexaura Medical Research Plexaura is a coral that has a large amount of prostaglandins making it a great natural source of a compound called Prostaglandin A2. Prostaglandins can be extracted from Plexaura using dichloromethane/methanol and a rotary evapo ...
'' species respectively. *The first ''Endozoicomonas'' sequenced sample was '' E. elysicola'', in 2013, being part of a project that focused on sequencing different strains of unknown microorganism at the time, and in 2014, its genome was updated and published together with the before unknown genomic sequences ''of E. montiporae'' and ''E. numazuensis.'' Then, '' E. atrinae'' was isolated from the intestine of the bivalve ''
Atrina pectinata ''Atrina pectinata'' is a species of bivalves belonging to the family Pinnidae. The species is found in the Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence ...
'' in 2014 as well. *After 2014, other species isolated and acknowledged were '' E. acroporae'' and ''E. ascidiicola'', the first one from a coral of the genus '' Acropora'' in Southern Taiwan and the latter from member of genus ''
Ascidiacea Ascidiacea, commonly known as the ascidians, tunicates (in part), and sea squirts (in part), is a polyphyletic class in the subphylum Tunicata of sac-like marine invertebrate filter feeders. Ascidians are characterized by a tough outer "tunic" ...
.'' At the moment, ten species are validly published under the ICNP.


Biology and Biochemistry


Genome

Despite the abundance of ''Endozoicomonas'' symbionts, only three complete ''Endozoicomonas'' genomes are publicly available (''E. elysicola'', ''E. montiporae'', and ''E. numazuensis)'', isolated from a
sea slug Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails (marine gastropod mollusks) that over evolutionary time ...
,
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...
, and a
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throug ...
, respectively. For their sequencing analyses, culture-independent methods of genome sequencing were used, including meta-genomic binning and single cell genomics. ''Endozoicomonas'' species have large genomes ranging from 4.049 Mb (''Endozoicomonas'' sp. AB1) to 6.69 Mb (''E. elysicola'' DSM22380).


Metabolism

Research led to the discovery that its genome is enriched with genes associated with transporter activity of carbon sugars, as well as cell secretion and
transposase A transposase is any of a class of enzymes capable of binding to the end of a transposon and catalysing its movement to another part of a genome, typically by a cut-and-paste mechanism or a replicative mechanism, in a process known as transpositio ...
activity, suggesting that these organisms have a potential role in the up-cycling of carbohydrates or the supply of proteins to their host. These skills can help them to quickly adapt to a new host or take advantage of a new niche. Although none of the ''Endozoicomonas'' genomes have genes for
fixing nitrogen Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmos ...
directly, some species have several forms of nitrate reductase, accounting for the conversion of nitrate to nitrite and of nitrite to ammonia, which could then be secreted. ''Endozoicomonas'' contain in their own genome for the assimilation of ammonia through the synthesis of glutamine and glutamate. They can also synthesize other
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...
s like alanine, aspartate, cysteine, glycine, homocysteine, homoserine, leucine, lysine, methionine, serine, and threonine, indicating strain-specific functions. The ''Endozoicomonas'' genus also plays an important role in the coral sulfur cycle. ''E. acroporae'' strains cannot only metabolize
dimethylsulfoniopropionate Dimethylsulfoniopropionate (DMSP), is an organosulfur compound with the formula (CH3)2S+CH2CH2COO−. This zwitterionic metabolite can be found in marine phytoplankton, seaweeds, and some species of terrestrial and aquatic vascular plants ...
(DMSP) to produce
dimethylsulfide Dimethyl sulfide (DMS) or methylthiomethane is an organosulfur compound with the formula (CH3)2S. Dimethyl sulfide is a flammable liquid that boils at and has a characteristic disagreeable odor. It is a component of the smell produced from cook ...
(DMS), but also use DMSP as a carbon source for growth and survival. Through several research done, the first DMSP-related operon in ''E. acroporae'' was also identified, which links DMSP metabolism to the central
carbon cycle The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth. Carbon is the main component of biological compounds as well as a major componen ...
. A high metabolic specificity is shown by ''Endozoicomonas'' samples isolated from the intertidal marine sponge O. ''papilla''. Indeed, the presence of gene clusters encoding for the lactate,
L-rhamnose Rhamnose (Rha, Rham) is a naturally occurring deoxy sugar. It can be classified as either a methyl-pentose or a 6-deoxy-hexose. Rhamnose predominantly occurs in nature in its L-form as L-rhamnose (6-deoxy-L-mannose). This is unusual, since most ...
metabolism, and phenylacetic acid (PA) degradation pathway indicates the probable ability of these microorganisms to utilize alternative carbon sources.


Ecology


Habitat

''Endozoicomonas'' are mutualistic organisms that have a symbiotic relationship with many marine animals. Found in all oceans of the world, they inhabit mostly in warm and mildly temperate waters located between the tropics, existing from the intertidal zone to the open ocean. Their most common association is the one shared with corals, especially with those found in shallow waters, but can also thrive in
deep-water coral The habitat of deep-water corals, also known as cold-water corals, extends to deeper, darker parts of the oceans than tropical corals, ranging from near the surface to the abyss, beyond where water temperatures may be as cold as . Deep-water co ...
s as well, locating themselves in the soft epithelial tissue of these. Also, they have been found to share this relationship with many other invertebrates such as sponges,
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ...
s, sea slugs, and some mollusks.


Role in the environment

The presence of ''Endozoicomonas'' in the marine ecosystem is associated with the overall coral health, serving as a marker of the general well-being of corals and the organisms that inhabit in coral reefs, as well as reducing the presence of pathogenic bacteria that may try to infect the coral. Other functions associated to ''Endozoicomonas'' relate to amino acid and vitamin synthesis, in the production of metabolites while contributing with nitrogen and sulfur cycles, and to transfer organic molecules which avidly helps in the nutrition of its host, yet their exact function and the way in which their presence affects all these organisms is still yet to be determined. During
coral bleaching Coral bleaching is the process when corals become white due to various stressors, such as changes in temperature, light, or nutrients. Bleaching occurs when coral polyps expel the zooxanthellae ( dinoflagellates that are commonly referred to as a ...
, ''Endozoicomonas'' populations remain present in the water in low amounts, indicating a certain level of resilience, and the absence of a healthy coral community leads to changes in the population amounts of these bacteria. Other environmental factors and stressors such as temperature changes, acidification of the ocean, and anthropogenic activities have a direct impact as well in the abundance of these microorganisms in their habitat. In contrast to their reputation as beneficial symbionts, their genome reveals potential mechanisms for bacterial adaptation and some pathogenic species are being discovered and described to be affecting fish larvae cultures, causing epitheliocystis and further leading to mass mortality.


References


Further reading

* * * Oceanospirillales Bacteria genera