Endozoicomonas
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''Endozoicomonas'' is a genus of
Gram-negative Gram-negative bacteria are bacteria that, unlike gram-positive bacteria, do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is that their cell envelope consists ...
, aerobic or facultatively anaerobic, chemoorganotrophic,
rod-shaped Bacterial cellular morphologies are the shapes that are characteristic of various types of bacteria and often key to their identification. Their direct examination under a light microscope enables the classification of these bacteria (and archae ...
, marine bacteria from the family of ''Endozoicomonadaceae''. ''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''. 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,'' 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'' 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'' 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.'' 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 biology, marine invertebrates with varying levels of resemblance to terrestrial Slug, slugs. Most creatures known as sea slugs are gastropods, i.e. they are Sea snail, sea snails (marine gastropod moll ...
,
coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
, and a
sponge Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a basal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and a ...
, 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 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 over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 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 The sulfur cycle is a biogeochemical cycle in which the sulfur moves between rocks, waterways and living systems. It is important in geology as it affects many minerals and in life because sulfur is an essential element (CHNOPS), being a consti ...
. ''E. acroporae'' strains cannot only metabolize dimethylsulfoniopropionate (DMSP) to produce dimethylsulfide (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 a part of the biogeochemical cycle where carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of Earth. Other major biogeochemical cycles include the nitrogen cycle and the water cycl ...
. 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 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 corals 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, tunicates, 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 loss of Symbiosis, symbiotic algae and Photosynthesis, photosynthetic pigments. This loss of pigment can be caused by various stressors, such as changes in water temperature, light, ...
, ''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