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Giant virus

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"Girus" redirects here. Not to be confused with Gyrus.

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]

Description

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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 in diameter)[4] that may be surrounded by a thick (approximately 100 nm) layer of filamentous protein fibers. The viruses have large, double-stranded DNA genomes (300 to >1000 kilobasepairs) that encode a large contingent of genes (of the order of 1000 genes).[3][5] The best characterized giant viruses are the phylogenetically related mimivirus and megavirus, which belong to the family Mimiviridae (aka Megaviridae), and are distinguished by their large capsid diameters.[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]

The genomes of many giant viruses encode many unusual genes that are not found in other viruses, including genes involved in glycolysis and the TCA cycle,[7] fermentation,[8] and the cytoskeleton.[9][10][11]

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.
Phylogeny of Nucleocytoviricota [12]

History

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The first giant viruses to be described were chloroviruses of the family Phycodnaviridae. These were discovered in 1981 by Russel H. Meints, James L. Van Etten, Daniel Kuczmarski, Kit Lee, and Barbara Ang. The first chlorovirus was initially called HVCV (Hydra viridis Chlorella virus) since it was first found to infect Chlorella-like algae.[13][14]

Other giant viruses that infected marine flagellates were described later. The first mimivirus (BV-PW1) was described in 1995,[15] but was not recognized as such until its sequenced genome was released as Cafeteria roenbergensis virus (CroV) in 2010.[16] Subsequently, the Giant Virus Acanthamoeba polyphaga Mimivirus was characterized[17] (which had been mistaken as a bacterium in 1993),[18] and then sequenced.[19] The term "girus" was coined to refer to the group in 2006.[20]

Genetics and evolution

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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 in the megavirus genome (including those in Mimiviridae) provide evidence that these large DNA viruses may have evolved from a shared cellular genome ancestor by means of genome reduction.[5]

The discovery and subsequent characterization of giant viruses has triggered debate on their evolutionary origins. The two main hypotheses are that they evolved from small viruses by picking up DNA from host organisms; or that they evolved from very complicated organisms via genome reduction, losing various functions including self-reproduction.[21] The possible complicated ancestral organism is also a topic of debate: by one proposal, it might represent a fourth domain of life,[5] but this has been largely discounted.[22][23][24]

Comparison of largest known giant viruses

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Largest giant viruses with complete sequenced genomes as of June 2018
Giant virus name Genome Length Genes Capsid diameter (nm) Hair cover Genbank #
Pandoravirus salinus[25] 2,473,870 2500 proteins (predicted) ~500 KC977571
Tupanvirus[26] 1,500,000 1276–1425 proteins ≥450+550[27] KY523104
MF405918[28]
Bodo saltans virus[29] 1,385,869 1227 proteins (predicted) ~300 yes (~40 nm) MF782455
Megavirus chilense[30] 1,259,197 1120 proteins (predicted) 440 yes (75 nm) JN258408
Mamavirus[31] 1,191,693 1023 proteins (predicted) 500 yes (120 nm) JF801956
Mimivirus[19][32] 1,181,549 979 proteins 39 non-coding 500 yes (120 nm) NC_014649
M4[33] (Mimivirus "bald" variant) 981,813 756 proteins (predicted) 390 No JN036606
Cafeteria roenbergensis virus[34] 617,453 (730 kb) 544 proteins (predicted) 300 No NC_014637

The whole list is in the Giant Virus Toplist created by the Giant Virus Finder software.[35] As of June 11, 2018, there were 183 listed.[36]

Specific common features among giant viruses
Giant virus name Aminoacyl-tRNA synthetase Octocoral-like 1MutS 2Stargate[37] Known virophage[38] Cytoplasmic virion factory Host
Megavirus chilensis 7 (Tyr, Arg, Met, Cys, Trp, Asn, Ile) yes yes no yes Acanthamoeba (Unikonta, Amoebozoa)
Mamavirus 4 (Tyr, Arg, Met, Cys) yes yes yes yes Acanthamoeba (Unikonta, Amoebozoa)
Mimivirus 4 (Tyr, Arg, Met, Cys) yes yes yes yes Acanthamoeba (Unikonta, Amoebozoa)
M4 (Mimivirus "bald" variant) 3 (Met, Cys, Arg) yes yes Resistant yes Acanthamoeba (Unikonta, Amoebozoa)
Cafeteria roenbergensis virus 1 (Ile) yes no yes yes Phagotrophic 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

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References

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