Mimivirus

Mimivirus
Virus classification
Group: Group I (dsDNA)
Family: Mimiviridae
Genus: Mimivirus
Species

Acanthamoeba polyphaga mimivirus
Acanthamoeba polyphaga moumouvirus
Mimivirus bombay
Courdo11 virus
LBA 111 virus
Samba virus
Shan virus
Terra1 virus
Terra2 virus

Mimivirus is a genus of viruses, in the family Mimiviridae. Amoeba serve as natural hosts.[1][2] This genus contains a single identified species named Acanthamoeba polyphaga mimivirus (APMV), which serves as its type species. It also refers to a group of phylogenetically related large viruses, designated usually "MimiN".[3]

In colloquial speech, APMV is more commonly referred to as just "mimivirus". Mimivirus, short for "mimicking microbe", is so called to reflect its large size and apparent Gram-staining properties.[4]

Mimivirus has a large and complex genome compared with most other viruses. Until October 2011, when a larger virus Megavirus chilensis was described, it had the largest capsid diameter of all known viruses.[5]

Discovery

APMV was discovered accidentally in 1992 within the amoeba Acanthamoeba polyphaga, after which it is named, during research into Legionellosis by researchers from Marseille and Leeds.[6] The virus was observed in a gram stain and mistakenly thought to be a gram-positive bacterium. As a consequence it was named Bradfordcoccus, after the district the amoeba was sourced from in Bradford, England. In 2003, researchers at the Université de la Méditerranée in Marseille, France published a paper in Science identifying the micro-organism as a virus. It was given the name "Mimivirus" (for "Mimicking microbe") as it resembles a bacterium on Gram staining.[7]

The same team that discovered the mimivirus later discovered a slightly larger virus, dubbed the mamavirus, and the Sputnik virophage that infects it.[8]

Classification

Mimivirus has been placed into a viral family by the International Committee on Taxonomy of Viruses as a member of the Mimiviridae,[9] and has been placed into Group I of the Baltimore classification system.[10]

Although not strictly a method of classification, Mimivirus joins a group of large viruses known as nucleocytoplasmic large DNA viruses (NCLDV). They are all large viruses which share both molecular characteristics and large genomes. The mimivirus genome also possesses 21 genes encoding homologs to proteins which are seen to be highly conserved in the majority of NCLDVs, and further work suggests that mimivirus is an early divergent of the general NCLDV group.[7]

Structure

A: AFM image of several surface fibers attached to a common central feature. B: AFM image of two detached surface fibers of Mimivirus. C: CryoEM image of a whole Mimivirus. D: AFM image of internal fibers of Mimivirus.

The mimivirus is the fourth-largest virus, preceded by the recently discovered Megavirus chilensis, Pandoravirus and Pithovirus. Mimivirus has a capsid diameter of 400 nm. Protein filaments measuring 100 nm project from the surface of the capsid, bringing the total length of the virus up to 600 nm. Variation in scientific literature renders these figures as highly approximate, with the "size" of the virion being casually listed as anywhere between 400 nm and 800 nm, depending on whether total length or capsid diameter is actually quoted.

Its capsid appears hexagonal under an electron microscope, therefore the capsid symmetry is icosahedral.[11] It does not appear to possess an outer viral envelope, suggesting that the virus does not exit the host cell by exocytosis.[12]

Mimivirus shares several morphological characteristics with all members of the NCLDV group of viruses. The condensed central core of the virion appears as a dark region under the electron microscope. The large genome of the virus resides within this area. An internal lipid layer surrounding the central core is present in all other NCLDV viruses, so this features may also be present in mimivirus.[11]

Several mRNA transcripts can be recovered from purified virions. Like other NCLDVs, transcripts for DNA polymerase, a capsid protein and a TFII-like transcription factor were found. However, three distinct aminoacyl tRNA synthetase enzyme transcripts and four unknown mRNA molecules specific to mimivirus were also found. These pre-packaged transcripts can be translated without viral gene expression and are likely to be necessary to Mimivirus for replication. Other DNA viruses, such as the Human cytomegalovirus and Herpes simplex virus type-1, also feature pre-packaged mRNA transcripts.[12]

Genus Structure Symmetry Capsid Genomic arrangement Genomic segmentation
Mimivirus Icosahedral T = 972–1141 or T = 1200 (h = 19 ± 1, k = 19 ± 1) Linear Monopartite

Genome

The mimivirus genome is a linear, double-stranded molecule of DNA with 1,181,404 base pairs in length.[13] This makes it one of the largest viral genomes known, outstripping the next-largest virus genome of the Cafeteria roenbergensis virus by about 450,000 base pairs. In addition, it is larger than at least 30 cellular clades.[14]

In addition to the large size of the genome, mimivirus possesses an estimated 979 protein-coding genes, far exceeding the minimum 4 genes required for viruses to exist (c.f. MS2 and Qβ viruses).[15] Analysis of its genome revealed the presence of genes not seen in any other viruses, including aminoacyl tRNA synthetases, and other genes previously thought only to be encoded by cellular organisms. Like other large DNA viruses, mimivirus contains several genes for sugar, lipid and amino acid metabolism, as well as some metabolic genes not found in any other virus.[12] Roughly 90% of the genome was of coding capacity, with the other 10% being "junk DNA".

Replication

The stages of mimivirus replication are not well known, but as a minimum it is known that mimivirus attaches to a chemical receptor on the surface of an amoeba cell and is taken into the cell. Once inside, an eclipse phase begins, in which the virus disappears and all appears normal within the cell. After about 4 hours small accumulations can be seen in areas of the cell. 8 hours after infection many mimivirus virions are clearly visible within the cell. The cell cytoplasm continues to fill with newly synthesised virions, and about 24 hours after initial infection the cell likely bursts open to release the new mimivirus virions.[12]

Little is known about the details of this replication cycle, most obviously attachment to the cell surface and entry, viral core release, DNA replication, transcription, translation, assembly and release of progeny virions. However, scientists have established the general overview given above using electron micrographs of infected cells. These micrographs show mimivirus capsid assembly in the nucleus, acquisition of an inner lipid membrane via budding from the nucleus, and particles similar to those found in many other viruses, including all NCLDV members. These particles are known in other viruses as viral factories and allow efficient viral assembly by modifying large areas of the host cell.

Genus Host details Tissue tropism Entry details Release details Replication site Assembly site Transmission
MimivirusZooplanktonNoneUnknownUnknownCytoplasmNucleusPassive diffusion

Pathogenicity

Mimivirus may be a causative agent of some forms of pneumonia; this is based mainly on indirect evidence in the form of antibodies to the virus discovered in pneumonia patients.[16] However, the classification of mimivirus as a pathogen is tenuous at present as there have been only a couple of papers published potentially linking mimivirus to actual cases of pneumonia. A significant fraction of pneumonia cases are of unknown etiology,[17] though a mimivirus has been isolated from a Tunisian woman suffering from pneumonia.[18]

Implications for defining "life"

Mimivirus shows many characteristics which place it at the boundary between living and non-living. It is as large as several bacterial species, such as Rickettsia conorii and Tropheryma whipplei, possesses a genomic size comparable to that of several bacteria, including those above, and codes for products previously not thought to be encoded by viruses (including a kind of collagen[19]). In addition, mimivirus has genes coding for nucleotide and amino acid synthesis, which even some small obligate intracellular bacteria lack. They do, however, lack any genes for ribosomal proteins, making mimivirus dependent on a host cell for protein translation and energy metabolism.[19]

Because its lineage is very old and could have emerged prior to cellular organisms,[20][21] Mimivirus has added to the debate over the origins of life. Some genes unique to Mimivirus, including those coding for the capsid, have been conserved in a variety of viruses which infect organisms from all domains. This has been used to suggest that Mimivirus is related to a type of DNA virus that emerged before cellular organisms and played a key role in the development of all life on Earth.[20] An alternative hypothesis is that there were three distinct types of DNA viruses that were involved in generating the three known domains of life—eukarya, archaea and bacteria.[21] It has been suggested that Mimivirus and similar kinds are remnants of a "fourth domain" of life, and that other giant virus may represent other ancient domains.[19]

Nevertheless, mimivirus does not exhibit the following characteristics, all of which are part of many conventional definitions of life:

See also

Wikispecies has information related to: Mimivirus

References

  1. ↑ "Viral Zone". ExPASy. Retrieved 15 June 2015.
  2. ↑ ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.
  3. ↑ Ghedin, E.; Claverie, J. (Aug 2005). "Mimivirus relatives in the Sargasso sea" (Free full text). Virology Journal. 2: 62. doi:10.1186/1743-422X-2-62. PMC 1215527Freely accessible. PMID 16105173.
  4. ↑ Wessner, D. R. (2010). "Discovery of the Giant Mimivirus". Nature Education. 3 (9): 61. Retrieved 2012-01-07.
  5. ↑ "World's biggest virus found in sea off Chile". London: Telegraph UK. 11 October 2011. Retrieved 11 November 2011.
  6. ↑ Richard Birtles, TJ Rowbotham, C Storey, TJ Marrie, Didier Raoult (29 Mar 1997). "Chlamydia-like obligate parasite of free-living amoebae". The Lancet. 349: 925–926. doi:10.1016/S0140-6736(05)62701-8. PMID 9093261.
  7. 1 2 Bernard La Scola; Stéphane Audic; Catherine Robert; Liang Jungang; Xavier de Lamballerie; Michel Drancourt; Richard Birtles; Jean-Michel Claverie; Didier Raoult. (2003). "A giant virus in amoebae". Science. 299 (5615): 2033. doi:10.1126/science.1081867. PMID 12663918.
  8. ↑ Nature: "'Virophage' suggests viruses are alive" by Helen Pearson 2008
  9. ↑ Claverie J-M (2010). Mahy W.J. and Van Regenmortel M. H. V., ed. Desk Encyclopedia of General Virology (1 ed.). Oxford: Academic Press. p. 189.
  10. ↑ Leppard, Keith; Nigel Dimmock; Easton, Andrew (2007). Introduction to Modern Virology (6 ed.). Blackwell Publishing Limited. pp. 469–470.
  11. 1 2 Xiao C., Kuznetsov Y. G., Sun S., Hafenstein S. L., Kostyuchenko V. A., Chipman P. R., Suzan-Monti M., Raoult D., McPherson A., Rossmann M. G. (2009-04-28). Sugden, Bill, ed. "Structural Studies of the Giant Mimivirus". PLoS Biol. 7 (4): e92. doi:10.1371/journal.pbio.1000092. PMC 2671561Freely accessible. PMID 19402750.
  12. 1 2 3 4 Suzan-Monti M., La Scola B., Raoult D. (2006). "Genomic and evolutionary aspects of Mimivirus". Virus Research. 117 (1): 145–155. doi:10.1016/j.virusres.2005.07.011. PMID 16181700.
  13. ↑ "Acanthamoeba polyphaga mimivirus, complete genome". NCBI.
  14. ↑ Claverie, Jean-Michel; et al. (2006). "Mimivirus and the emerging concept of 'giant' virus". Virus Research. 117 (1): 133–144. doi:10.1016/j.virusres.2006.01.008. PMID 16469402.
  15. ↑ Prescott, Lansing M. (1993). 2nd edition, ed. Microbiology. Dubuque, IA: Wm. C. Brown Publishers. ISBN 0-697-01372-3.
  16. ↑ La Scola B, Marrie T, Auffray J, Raoult D (2005). "Mimivirus in pneumonia patients". Emerg Infect Dis. 11 (3): 449–52. doi:10.3201/eid1103.040538. PMC 3298252Freely accessible. PMID 15757563.
  17. ↑ Marrie TJ, Durant H, Yates L (1989). "Community-Acquired Pneumonia Requiring Hospitalization: 5-Year Prospective Study". Reviews of Infectious Diseases. 11 (4): 586–99. doi:10.1093/clinids/11.4.586. PMID 2772465.
  18. ↑ Saadi, H, Pagnier, I; Colson, P; Cherif, JK; Beji, M; Boughalmi, M; Azza, S; Armstrong, N; Robert, C; Fournous, G; La Scola, B; Raoult, D (Jun 21, 2013). "First Isolation of Mimivirus in a Patient With Pneumonia.". Clinical Infectious Diseases. 57 (4): e127–34. doi:10.1093/cid/cit354. PMID 23709652.
  19. 1 2 3 Garry Hamilton (Jan 23, 2016). "How giant viruses could rewrite the story of life on Earth". New Scientist.
  20. 1 2 Siebert, Charles (2006-03-15). "Unintelligent Design". Discover Magazine.
  21. 1 2 Forterre, Patrick (2006). "Three RNA cells for ribosomal lineages and three DNA viruses to replicate their genomes: A hypothesis for the origin of cellular domain". PNAS. 103 (10): 3669–3674. doi:10.1073/pnas.0510333103. PMC 1450140Freely accessible. PMID 16505372.

Further reading

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