Zamilon virophage

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Mimivirus-dependent virus Zamilon
Giant virus Megavirus with its associated virophage Zamilon.png
Two Zamilon virophages (inset) to scale with their associated giant virus Megavirus [1]
Virus classification OOjs UI icon edit-ltr.svg
(unranked): Virus
Realm: Varidnaviria
Kingdom: Bamfordvirae
Phylum: Preplasmiviricota
Class: Maveriviricetes
Order: Priklausovirales
Family: Lavidaviridae
Genus: Sputnikvirus
Species:
Mimivirus-dependent virus Zamilon

Mimivirus-dependent virus Zamilon, or Zamilon, is a virophage, a group of small DNA viruses that infect protists and require a helper virus to replicate; they are a type of satellite virus. [2] Discovered in 2013 in Tunisia, infecting Acanthamoeba polyphaga amoebae, Zamilon most closely resembles Sputnik, the first virophage to be discovered. The name is Arabic for "the neighbour". [3] Its spherical particle is 50–60 nm in diameter, and contains a circular double-stranded DNA genome of around 17 kb, which is predicted to encode 20 polypeptides. A related strain, Zamilon 2, has been identified in North America.

Contents

All known virophages are associated with helpers in the giant DNA virus family Mimiviridae . Zamilon is restricted in its range of helper viruses; it can be supported by viruses from Mimivirus -like Mimiviridae lineages B and C, but not from lineage A. This appears to be a consequence of a rudimentary immune system of the helper virus, termed MIMIVIRE (mimivirus virophage resistance element), akin to the CRISPR-Cas pathway. [4] Unlike the Sputnik virophage, Zamilon does not appear to impair the replication of its helper virus.

Zamilon was discovered in 2013, in Acanthamoeba polyphaga amoebae co-infected with the giant virus Mont1, isolated from a Tunisian soil sample. [2] [3] [5] As of 2015, Zamilon is one of three virophages to have been isolated physically, the others being Sputnik and Mavirus; several other virophage DNAs have been discovered using metagenomics but have not been characterised physically. [2] [6] A related strain, named Zamilon 2, was discovered by metagenomic analysis of a North American poplar wood chip bioreactor in 2015. [7] Another virophage, Rio Negro, is also closely related to Sputnik. [2]

Taxonomy

Zamilon virophage has been classified by the International Committee on Taxonomy of Viruses into the species Mimivirus-dependent virus Zamilon within the genus Sputnikvirus in the family Lavidaviridae . [2]

Virology

A Zamilon virion is spherical with a diameter of 50–60 nm, and is similar in appearance to those of Sputnik and Mavirus. [3] Its circular double-stranded DNA genome is 17,276  base pairs in length. [2] [3] Virophages typically have particles whose diameter is in the range 40–80 nm, with genomes in the range 17–30 kb. [2] The virophage most closely related to Zamilon is Sputnik, with which it shares 76% sequence identity, although part of the Zamilon sequence is reversed compared with Sputnik. [2] [3] Zamilon's DNA is rich in adenine and thymine bases; the proportion of guanine and cytosine bases is 29.7%. [3]

Open reading frames

The Zamilon genome is predicted to contain 20 open reading frames (ORFs), of between 222 bases and 2337 bases in length. Of the 20 predicted products, 15 are similar to those of Sputnik, and three are similar to Mimiviridae : two to Megavirus chilensis and another to the transpoviron of Moumouvirus monve. One Zamilon ORF additionally shows some similarity to Mavirus, and another to Organic Lake virophage and a Phaeocystis globosa virophage, which are both associated with algae rather than amoebae. The remaining two predicted products show limited similarity to other known proteins. Putative functions of the products include transposase, helicase, integrase, cysteine protease, DNA primasepolymerase and DNA-packaging ATPase enzymes, major and minor capsid proteins, a structural protein and a collagen-like protein. [3] ORF6 is very similar to the Sputnik major capsid protein, which has a double "jelly-roll" fold. [3] [8]

ORF [lower-alpha 1] Predicted size
(amino acids)
Related productSimilarity
(%)
Predicted function
1111none
273none
3135Megavirus chilensis mg3 gene product67
4221Sputnik virophage 2 putative IS3 family transposase A protein40putative transposase
5376Sputnik virophage 2 minor virion protein66minor virion protein
6609Sputnik virophage capsid protein V2086capsid protein
7442Sputnik virophage V2170
881Moumouvirus monve hypothetical protein tv_L872
9778Sputnik virophage V1367putative helicase
10168Sputnik virophage V1153
11247Sputnik virophage V1058putative integrase
12175Sputnik virophage V977
13184Sputnik virophage V871structural protein
14241Sputnik virophage V780
15305Sputnik virophage V675collagen-like protein
16121Sputnik virophage V559
17133Sputnik virophage V455
18245Sputnik virophage V381DNA-packaging ATPase
19147Megavirus chilensis mg664 gene product50
20147Sputnik virophage V160

Life cycle and helper virus

Like all other virophages, Zamilon replicates in the cytoplasm, within the virus factory of its helper, which acts as its host. [3] Zamilon was first isolated in association with Mont1, a Mimivirus-like Mimiviridae classified in lineage C by its polymerase B gene sequence. The virophage has subsequently been shown to be capable of replicating in association with Moumouvirus and Monve, two Mimiviridae from lineage B, as well as with Terra1 and Courdo11 from lineage C; however, it cannot replicate in association with either Mimivirus or Mamavirus , classified as lineage A. [3] [9] This is unlike Sputnik, which can replicate in association with any Mimivirus-like member of Mimiviridae. [3]

Zamilon does not appear to inhibit the ability of its helper virus to replicate significantly, nor to lyse its host amoebae cells. Although the helper virus formed a high proportion of abnormal virions in the presence of Zamilon, these were also observed at a comparable level in the virophage's absence. [3] This is again unlike Sputnik, which reduces its helper virus's infectivity, inhibits its lysis of amoeba, and is associated with the generation of an increased proportion of abnormal Mimiviridae virions. [2] [3] Bernard La Scola and colleagues, who isolated both Sputnik and Zamilon, state that, if confirmed, this "would question the concept of virophage", which has been considered to be differentiated from most other satellite viruses in having a deleterious effect on its helper virus. [3]

Related Research Articles

<i>Mimivirus</i> Genus of viruses

Mimivirus is a genus of giant viruses, in the family Mimiviridae. Amoeba serve as their natural hosts. This genus contains a single identified species named Acanthamoeba polyphaga mimivirus (APMV). It also refers to a group of phylogenetically related large viruses.

<span class="mw-page-title-main">Satellite (biology)</span> Subviral agent which depends on a helper virus for its replication

A satellite is a subviral agent that depends on the coinfection of a host cell with a helper virus for its replication. Satellites can be divided into two major classes: satellite viruses and satellite nucleic acids. Satellite viruses, which are most commonly associated with plants, are also found in mammals, arthropods, and bacteria. They encode structural proteins to enclose their genetic material, which are therefore distinct from the structural proteins of their helper viruses. Satellite nucleic acids, in contrast, do not encode their own structural proteins, but instead are encapsulated by proteins encoded by their helper viruses. The genomes of satellites range upward from 359 nucleotides in length for satellite tobacco ringspot virus RNA (STobRV).

<span class="mw-page-title-main">Viroplasm</span>

A viroplasm, sometimes called "virus factory" or "virus inclusion", is an inclusion body in a cell where viral replication and assembly occurs. They may be thought of as viral factories in the cell. There are many viroplasms in one infected cell, where they appear dense to electron microscopy. Very little is understood about the mechanism of viroplasm formation.

<span class="mw-page-title-main">Mobilome</span>

The mobilome is the entire set of mobile genetic elements in a genome. Mobilomes are found in eukaryotes, prokaryotes, and viruses. The compositions of mobilomes differ among lineages of life, with transposable elements being the major mobile elements in eukaryotes, and plasmids and prophages being the major types in prokaryotes. Virophages contribute to the viral mobilome.

<span class="mw-page-title-main">Virophage</span> Viral parasites of giant viruses

Virophages are small, double-stranded DNA viral phages that require the co-infection of another virus. The co-infecting viruses are typically giant viruses. Virophages rely on the viral replication factory of the co-infecting giant virus for their own replication. One of the characteristics of virophages is that they have a parasitic relationship with the co-infecting virus. Their dependence upon the giant virus for replication often results in the deactivation of the giant viruses. The virophage may improve the recovery and survival of the host organism.

<span class="mw-page-title-main">Sputnik virophage</span>

Mimivirus-dependent virus Sputnik is a subviral agent that reproduces in amoeba cells that are already infected by a certain helper virus; Sputnik uses the helper virus's machinery for reproduction and inhibits replication of the helper virus. It is known as a virophage, in analogy to the term bacteriophage.

<i>Mimiviridae</i> Family of viruses

Mimiviridae is a family of viruses. Amoeba and other protists serve as natural hosts. The family is divided in up to 4 subfamilies. Viruses in this family belong to the nucleocytoplasmic large DNA virus clade (NCLDV), also referred to as giant viruses.

Mamavirus is a large and complex virus in the Group I family Mimiviridae. The virus is exceptionally large, and larger than many bacteria. Mamavirus and other mimiviridae belong to nucleocytoplasmic large DNA virus (NCLDVs) family. Mamavirus can be compared to the similar complex virus mimivirus; mamavirus was so named because it is similar to but larger than mimivirus.

Marseillevirus is a genus of viruses, in the family Marseilleviridae. There are two species in this genus. It is the prototype of a family of nucleocytoplasmic large DNA viruses (NCLDV) of eukaryotes. It was isolated from amoeba.

<i>Cafeteria roenbergensis virus</i> Species of virus

Cafeteria roenbergensis virus (CroV) is a giant virus that infects the marine bicosoecid flagellate Cafeteria roenbergensis, a member of the microzooplankton community.

A giant virus, sometimes referred to as a girus, is a very large virus, some of which are larger than typical bacteria. All known giant viruses belong to the phylum Nucleocytoviricota.

<i>Marseilleviridae</i> Family of viruses

Marseilleviridae is a family of viruses first named in 2012. The genomes of these viruses are double-stranded DNA. Amoeba are often hosts, but there is evidence that they are found in humans as well. The family contains one genus and four species, two of which are unassigned to a genus. It is a member of the nucleocytoplasmic large DNA viruses clade.

<span class="mw-page-title-main">Megavirus</span> Genus of viruses

Megavirus is a viral genus containing a single identified species named Megavirus chilense, phylogenetically related to Acanthamoeba polyphaga Mimivirus (APMV). In colloquial speech, Megavirus chilensis is more commonly referred to as just “Megavirus”. Until the discovery of pandoraviruses in 2013, it had the largest capsid diameter of all known viruses, as well as the largest and most complex genome among all known viruses.

<i>Pandoravirus</i> Genus of giant virus possessing a large double-stranded DNA genome

Pandoravirus is a genus of giant virus, first discovered in 2013. It is the second largest in physical size of any known viral genus. Pandoraviruses have double stranded DNA genomes, with the largest genome size of any known viral genus.

<i>Pithovirus</i> Genus of viruses

Pithovirus, first described in a 2014 paper, is a genus of giant virus known from two species, Pithovirus sibericum, which infects amoebas and Pithovirus massiliensis. It is a double-stranded DNA virus and is a member of the nucleocytoplasmic large DNA viruses clade. The 2014 discovery was made when a viable specimen was found in a 30,000-year-old ice core harvested from permafrost in Siberia, Russia.

<i>Faustovirus</i> Genus of viruses

Faustovirus is a genus of giant virus which infects amoebae associated with humans. The virus was first isolated in 2015 and shown to be around 0.2 micrometers in diameter with a double stranded DNA genome of 466 kilobases predicted to encode 451 proteins. Although classified as a nucleocytoplasmic large DNA virus (NCLDV), faustoviruses share less than a quarter of their genes with other NCLDVs; however, ~46% are homologous to bacterial genes and the remainder are orphan genes (ORFans). Specifically, the gene encoding the major capsid protein (MCP) of faustovirus is different than that of its most closely related giant virus, asfivirus, as well as other NCLDVs. In asfivirus, the gene encoding MCP is a single genomic fragment of ~2000 base pairs (bp), however, in faustovirus the MCP is encoded by 13 exons separated by 12 large introns. The exons have a mean length of 149 bp and the introns have a mean length of 1,273 bp. The presence of introns in faustovirus genes is highly unusual for viruses.

<i>Tupanvirus</i> Proposed genus of viruses

Tupanvirus is a genus of viruses first described in 2018. The genus is composed of two species of virus that are in the giant virus group. Researchers discovered the first isolate in 2012 from deep water sediment samples taken at 3000m depth off the coast of Brazil. The second isolate was collected from a soda lake in Southern Nhecolândia, Brazil in 2014. They are named after Tupã (Tupan), a Guaraní thunder god, and the places they were found. These are the first viruses reported to possess genes for amino-acyl tRNA synthetases for all 20 standard amino acids.

<span class="mw-page-title-main">Chrysochromulina ericina virus</span> Giant virus

Chrysochromulina ericina virus 01B, or simply Chrysochromulina ericina virus (CeV) is a giant virus in the family Mimiviridae infecting Haptolina ericina, a marine microalgae member of the Haptophyta. CeV is a dsDNA virus.

<i>Varidnaviria</i> Realm of viruses

Varidnaviria is a realm of viruses that includes all DNA viruses that encode major capsid proteins that contain a vertical jelly roll fold. The major capsid proteins (MCP) form into pseudohexameric subunits of the viral capsid, which stores the viral deoxyribonucleic acid (DNA), and are perpendicular, or vertical, to the surface of the capsid. Apart from this, viruses in the realm also share many other characteristics, such as minor capsid proteins (mCP) with the vertical jelly roll fold, an ATPase that packages viral DNA into the capsid, and a DNA polymerase that replicates the viral genome.

Nucleocytoviricota is a phylum of viruses. Members of the phylum are also known as the nucleocytoplasmic large DNA viruses (NCLDV), which serves as the basis of the name of the phylum with the suffix -viricota for virus phylum. These viruses are referred to as nucleocytoplasmic because they are often able to replicate in both the host's cell nucleus and cytoplasm.

References

  1. All data from Gaia et al. [3]
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