Olavius algarvensis
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''Olavius algarvensis'' is a species of gutless
oligochaete worm Oligochaeta () is a subclass of animals in the phylum Annelida, which is made up of many types of aquatic and terrestrial worms, including all of the various earthworms. Specifically, oligochaetes comprise the terrestrial megadrile earthworms ...
in the family
Tubificidae The Naididae (including the former family Tubificidae) are a family of clitellate oligochaete worms like the sludge worm, '' Tubifex tubifex''. They are key components of the benthic communities of many freshwater and marine ecosystems. In fres ...
which depends on
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
bacteria for its nutrition.


Habitats and research

''Olavius algarvensis'' lives in coastal sediments in the Mediterranean. It was first described from the
Algarve The Algarve (, , ; from ) is the southernmost NUTS II region of continental Portugal. It has an area of with 467,495 permanent inhabitants and incorporates 16 municipalities ( ''concelhos'' or ''municípios'' in Portuguese). The region has it ...
Coast of Portugal, but has also been found elsewhere, e.g. off the Italian island
Elba Elba ( it, isola d'Elba, ; la, Ilva) is a Mediterranean island in Tuscany, Italy, from the coastal town of Piombino on the Italian mainland, and the largest island of the Tuscan Archipelago. It is also part of the Arcipelago Toscano National ...
, where it co-occurs with another species, '' O. ilvae.'' It was the first species of ''Olavius'' described from the East Atlantic coast; previously the genus was only known from the Caribbean.


Description

''Olavius algarvensis'' is 12–25 mm long, about 0.25 mm wide, and has between 100 and 150 segments. Like all other species in the genus ''Olavius'', this species has no digestive tract. Instead, the body cavity contains the
ventral nerve cord The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, in ...
(inside a muscular sheath) and two blood vessels which are surrounded by a "fluffy" layer of chloragocytic cells. They are distinguished from other species of ''Olavius'' by having round, flap-like external male papillae that cover the two ventral invaginations of the body wall which contain the male pores (in segment XI), and having small atria that are perpendicular rather than parallel to the body axis. The symbiotic bacteria are located between the cuticle and epidermis, and also in vacuoles within epidermal cells, which often show signs of lysis. The bacteria are absent from the anterior part of the worm and the
pygidium The pygidium (plural pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compo ...
, but are found from segment VII or VIII onwards.


Symbiosis with bacteria

Oligochaete worms without any
mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
, gut, or nephridial excretory system were first discovered in the 1970s-1980s near
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. They were later found to contain symbiotic
chemosynthetic In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydro ...
bacteria which serve as their primary food source. ''O. algarvensis'' is the species where this symbiosis has been studied in the most detail. There are five different species of bacterial
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
in ''O. algarvensis'', which are located under the cuticle of the worm: two sulfide-oxidizing
Gammaproteobacteria Gammaproteobacteria is a class of bacteria in the phylum Pseudomonadota (synonym Proteobacteria). It contains about 250 genera, which makes it the most genera-rich taxon of the Prokaryotes. Several medically, ecologically, and scientifically imp ...
, two sulfate-reducing
Deltaproteobacteria The Myxococcota are a phylum of bacteria known as the fruiting gliding bacteria. All species of this group are Gram-negative. They are predominantly aerobic genera that release myxospores in unfavorable environments. Phylogeny The currently acce ...
, and one
spirochaete A spirochaete () or spirochete is a member of the phylum Spirochaetota (), (synonym Spirochaetes) which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or s ...
. The sulfide-oxidizers gain energy from oxidation of hydrogen sulfide, and fix carbon dioxide via the
Calvin cycle The Calvin cycle, light-independent reactions, bio synthetic phase, dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis is a series of chemical reactions that convert carbon dioxide and hydrogen-carrier compounds into ...
. The sulfate-reducers are anaerobes that can reduce sulfate into sulfide, which is consumed by the sulfide-oxidizers. The metabolism of the spirochaete is unknown. Other species of ''Olavius'' are also known to have similar symbioses with both sulfide-oxidizing and sulfate-reducing bacteria in the same worm. The primary sulfur-oxidizing symbiont, known as "Gamma1", is closely related to the primary symbionts of other species of gutless oligochaetes in the
Phallodrilinae Phallodrilinae is a subfamily of clitellate oligochaete worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in s ...
, and also to the symbionts of nematodes in the subfamily Stilbonematinae. In addition to hydrogen sulfide, the symbiotic bacteria also allow the worm to use
hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-toxic, an ...
and
carbon monoxide Carbon monoxide (chemical formula CO) is a colorless, poisonous, odorless, tasteless, flammable gas that is slightly less dense than air. Carbon monoxide consists of one carbon atom and one oxygen atom connected by a triple bond. It is the simple ...
as energy sources, and to metabolise organic compounds like
malate Malic acid is an organic compound with the molecular formula . It is a dicarboxylic acid that is made by all living organisms, contributes to the sour taste of fruits, and is used as a food additive. Malic acid has two stereoisomeric forms (L ...
and
acetate An acetate is a salt (chemistry), salt formed by the combination of acetic acid with a base (e.g. Alkali metal, alkaline, Alkaline earth metal, earthy, Transition metal, metallic, nonmetallic or radical Radical (chemistry), base). "Acetate" als ...
. These abilities were first discovered by sequencing the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
s and
proteome The proteome is the entire set of proteins that is, or can be, expressed by a genome, cell, tissue, or organism at a certain time. It is the set of expressed proteins in a given type of cell or organism, at a given time, under defined conditions. ...
s of the bacteria. The symbiotic bacteria which live with ''O. algarvensis'' have other unique properties. One of the Deltaproteobacteria symbionts, called "Delta-1", is able to produce numerous seleno- and pyrroproteins, which contain the amino acids
selenocysteine Selenocysteine (symbol Sec or U, in older publications also as Se-Cys) is the 21st proteinogenic amino acid. Selenoproteins contain selenocysteine residues. Selenocysteine is an analogue of the more common cysteine with selenium in place of the s ...
and
pyrrolysine Pyrrolysine (symbol Pyl or O; encoded by the 'amber' stop codon UAG) is an α-amino acid that is used in the biosynthesis of proteins in some methanogenic archaea and bacteria; it is not present in humans. It contains an α-amino group (which is ...
that are sometimes called the 21st and 22nd
proteinogenic amino acid Proteinogenic amino acids are amino acids that are incorporated biosynthetically into proteins during translation. The word "proteinogenic" means "protein creating". Throughout known life, there are 22 genetically encoded (proteinogenic) amino aci ...
s. This bacterium has the largest known proteome that has seleno- and pyrroproteins. The symbionts also express the most
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 transposition ...
s of any known bacteria.


References

{{Taxonbar, from=Q1554464 Tubificina Animals described in 1998 Chemosynthetic symbiosis Taxa named by Christer Erséus