Epixenosomes, also known as

Candidatus In prokaryote nomenclature, ''Candidatus'' (abbreviated ''Ca.''; Latin for "candidate of Roman office") is used to name prokaryotic taxa that are well characterized but yet- uncultured. Contemporary sequencing approaches, such as 16S ribosomal R ...

''Epixenosoma'', are a genus of

bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

in the phylum

Verrucomicrobiota Verrucomicrobiota is a phylum of Gram-negative bacteria that contains only a few described species. The species identified have been isolated from fresh water, marine and soil environments and human faeces. A number of as-yet uncultivated species ...

that form a

symbiosis Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...

with marine

ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...

s of the genus '' Euplotidium'', where they help to defend their ciliate hosts against predators. It is a

monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...

genus, containing only the species Ca. ''Epixenosoma ejectans''.

Description

Epixenosomes possess two distinct developmental phases linked to the host cell cycle. Stage I epixenosomes are spherical, 0.5 μm in diameter and are surrounded by two

membranes A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. B ...

. They divide by direct binary

fission Fission, a splitting of something into two or more parts, may refer to: * Fission (biology), the division of a single entity into two or more parts and the regeneration of those parts into separate entities resembling the original * Nuclear fissio ...

. Stage II epixenosomes are egg-shaped (2.2 μm in length and 1 μm in width) and show complex organization with different

cytoplasm The cytoplasm describes all the material within a eukaryotic or prokaryotic cell, enclosed by the cell membrane, including the organelles and excluding the nucleus in eukaryotic cells. The material inside the nucleus of a eukaryotic cell a ...

ic compartments, more complicated than the majority of

prokaryotic A prokaryote (; less commonly spelled procaryote) is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Greek (), meaning 'before', and (), meaning 'nut' ...

organisms. They contain an extrusive apparatus within a

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...

aceous matrix, although apparently not membrane-bound, which differs from the remaining

. A functional cell compartmentalization has also been evidenced. Their

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

position was originally unclear, because they appeared to have both prokaryote-like traits, such as binary fission, and

eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

-like traits, such as intracellular membranes. However,

molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

showed that they are actually bacteria from the phylum

Defensive symbiosis with ''Euplotidium''

Epixenosomes live on the

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...

surface of their hosts, marine ciliates in the genus ''Euplotidium''. The name "epixenosomes" comes from the ancient Greek , meaning "external alien body", referring to their extracellular position on the host. The extrusive apparatus ejects its contents when triggered; this process helps to defend the ciliate host against

predators Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...

. Although ''Euplotidium'' can grow and reproduce without epixenosomes, those with epixenosomes have much higher survival when exposed to predators such as the ciliate '' Litonotus''. External signals of unknown origin are detected by the membrane receptors located at the top of the organism. The consequent activation of the

adenylate cyclase Adenylate cyclase (EC 4.6.1.1, also commonly known as adenyl cyclase and adenylyl cyclase, abbreviated AC) is an enzyme with systematic name ATP diphosphate-lyase (cyclizing; 3′,5′-cyclic-AMP-forming). It catalyzes the following reaction: :A ...

cAMP Camp may refer to: Areas of confinement, imprisonment, or for execution * Concentration camp, an internment camp for political prisoners or politically targeted demographics, such as members of national or minority ethnic groups * Extermination ...

system triggers the ejection of the extrusive apparatus, with the formation of a hollow tube, about 40 μm long, terminating in a head mainly consisting of the apical portion of the epixenosome (the region containing

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

). The extrusive apparatus is surrounded by a basket of tubules. Experiments with antitubulin drugs and immunocytochemical analyses at the optical and electron microscopical level suggested that these tubules consist of

tubulin Tubulin in molecular biology can refer either to the tubulin protein superfamily of globular proteins, or one of the member proteins of that superfamily. α- and β-tubulins polymerize into microtubules, a major component of the eukaryotic cytosk ...

, which is a

eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

. Although some free-living Verrucomicrobiota are reported to have tubulin genes, they have not been found to have tubular structures like those of epixenosomes.

References

{{Taxonbar, from=Q3730697, from2=Q51536694 Candidatus taxa Verrucomicrobiota Cell biology Symbiosis Monotypic bacteria genera

Description

Defensive symbiosis with ''Euplotidium''

See also

References