Giant virus

A giant virus, sometimes referred to as a girus, is a very large virus, some of which are larger than typical bacteria.[1][2] All known giant viruses belong to the phylum Nucleocytoviricota.[3]

"Girus" redirects here. It is not to be confused with Gyrus.
Megaviricetes
Mimivirus
Virus classification
(unranked): Virus
Realm: Varidnaviria
Kingdom: Bamfordvirae
Phylum: Nucleocytoviricota
Class: Megaviricetes

Description

While the exact criteria as defined in the scientific literature vary, giant viruses are generally described as viruses having large, pseudo-icosahedral capsids (200 to 400 nanometers)[4] that may be surrounded by a thick (approximately 100 nm) layer of filamentous protein fibers. The viruses' large, double-stranded DNA genomes (300 to 1000 kilobasepairs or larger) encode a large contingent of genes (of the order of 1000 genes).[3][5] While few giant viruses have been characterized in detail, the most notable examples are the phylogenetically related mimivirus and megavirus—both belonging to the Mimiviridae (aka Megaviridae) family, due to their having the largest capsid diameters of all known viruses.[3][5]Giant viruses from the deep ocean, terrestrial sources, and human patients contain genes encoding cytochrome P450 (CYP; P450) enzymes. The origin of these P450 genes in giant viruses remains unknown but may have been acquired from an ancient host.[6]

Cryo-EM images of the giant viruses CroV and APMV. (A) Cryo-electron micrograph of four CroV particles. (B) Single CroV particle with concave core depression (white arrow). (C) Single APMV particle. Scale bars in (A–C) represent 2,000 Å.

History

The first giant virus, then named BV-PW1, was isolated and described in 1995,[7] but was not recognized as such until its sequenced genome was released as Cafeteria roenbergensis virus (CroV) in 2010.[8] Subsequently, the Giant Virus Acanthamoeba polyphaga Mimivirus was characterized,[9] (which had been mistaken as a bacterium in 1993),[10] and then sequenced.[11] The term "girus" was coined to refer to the group in 2006.[12]

Genetics and evolution

The genomes of giant viruses are the largest known for viruses, and contain genes that encode for important elements of translation machinery, a characteristic that had previously been believed to be indicative of cellular organisms. These genes include multiple genes encoding a number of aminoacyl tRNA synthetases, enzymes that catalyze the esterification of specific amino acids or their precursors to their corresponding cognate tRNAs to form an aminoacyl tRNA that is then used during translation.[5] The presence of four aminoacyl tRNA synthetase encoding genes in mimivirus and mamavirus genomes, both species within the Mimiviridae family, as well as the discovery of seven aminoacyl tRNA synthetase genes, including the four genes present in Mimiviridae, in the megavirus genome provide evidence for a possible scenario in which these large DNA viruses evolved from a shared ancestral cellular genome by means of genome reduction.[5]

Their discovery and subsequent characterization has triggered some debate concerning the evolutionary origins of giant viruses. The two main hypotheses for their origin are that either they evolved from small viruses, picking up DNA from host organisms, or that they evolved from very complicated organisms into the current form which is not self-sufficient for reproduction.[13] What sort of complicated organism giant viruses might have diverged from is also a topic of debate. One proposal is that the origin point actually represents a fourth domain of life,[5] but this has been largely discounted.[14][15]

Comparison of largest known giant viruses
Largest giant viruses with complete sequenced genomes as of March 2015
Giant virus nameGenome LengthGenesCapsid diameter (nm)Hair coverGenbank #
Bodo saltans virus[16]1,385,8691227 proteins (predicted)~300yes (~40 nm)MF782455
Megavirus chilensis[17]1,259,1971120 proteins (predicted)440yes (75 nm)JN258408
Mamavirus[18]1,191,6931023 proteins (predicted)500yes (120 nm)JF801956
Mimivirus[11][19]1,181,549979 proteins 39 non-coding500yes (120 nm)NC_014649
M4[20] (Mimivirus "bald" variant)981,813756 proteins (predicted)390NoJN036606
Tupanvirus[21]1,500,0001276–1425 proteins≥450+550[22]KY523104
MF405918[23]
Cafeteria roenbergensis virus[24]617,453 (730 kb)544 proteins (predicted)300NoNC_014637

The whole list is in the Giant Virus Toplist created by the Giant Virus Finder software.[25]

Specific common features among giant viruses
Giant virus nameAminoacyl-tRNA synthetaseOctocoral-like 1MutS2Stargate[26]Known virophage[27]Cytoplasmic virion factoryHost
Megavirus chilensis7 (Tyr, Arg, Met, Cys, Trp, Asn, Ile)yesyesnoyesAcanthamoeba (Unikonta, Amoebozoa)
Mamavirus4 (Tyr, Arg, Met, Cys)yesyesyesyesAcanthamoeba (Unikonta, Amoebozoa)
Mimivirus4 (Tyr, Arg, Met, Cys)yesyesyesyesAcanthamoeba (Unikonta, Amoebozoa)
M4 (Mimivirus "bald" variant)3 (Met, Cys, Arg)yesyesResistantyesAcanthamoeba (Unikonta, Amoebozoa)
Cafeteria roenbergensis virus1 (Ile)yesnoyesyesPhagotrophic protozoan (Heterokonta, Stramenopiles)

1Mutator S (MutS) and its homologs are a family of DNA mismatch repair proteins involved in the mismatch repair system that acts to correct point mutations or small insertion/deletion loops produced during DNA replication, increasing the fidelity of replication. 2A stargate is a five-pronged star structure present on the viral capsid forming the portal through which the internal core of the particle is delivered to the host's cytoplasm.

See also

Notes

    References
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    22. head and tail, respectively
    23. soda lake and deep ocean species of Tupanvirues, respectively
    24. Fischer, M. G.; Allen, M. J.; Wilson, W. H.; Suttle, C. A. (2010). "Giant virus with a remarkable complement of genes infects marine zooplankton" (PDF). Proceedings of the National Academy of Sciences. 107 (45): 19508–13. Bibcode:2010PNAS..10719508F. doi:10.1073/pnas.1007615107. PMC 2984142. PMID 20974979.
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