Giant virus

A giant virus, also known as a girus, is a very large virus, some of which are larger than typical bacteria.[1][2] They have extremely large genomes compared to other viruses and contain many unique genes not found in life forms. All known giant viruses belong to the phylum Nucleocytoviricota (nucleocytoplasmic large DNA viruses, NCLDVs); they are the larger members of this group.[3]

Not to be confused with Gyrus.

Giant virus
Mimivirus
Virus classification
Group:
Group I (dsDNA)
Phylum:
Nucleocytoviricota

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) 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][4] 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][4]

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.

Viral replication in giant viruses occurs within large circular virus factories located within the cytoplasm of the infected host cell. This is similar to the replication mechanism used by Poxviridae, though whether this mechanism is employed by all giant viruses or only mimivirus and the related mamavirus has yet to be determined.[4] These virion replication factories are themselves subject to infection by the virophage satellite viruses, which inhibit or impair the reproductive capabilities of the complementary virus.

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.[5]

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.[4] 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.[4]

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.[6] 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,[4] but this has been largely discounted.[7][8]

Comparison of largest known giant viruses
Table 1 - Largest giant viruses with complete sequenced genomes, as of March 2015
Giant virus nameGenome LengthGenesCapsid diameter (nm)Hair coverGenbank #
Bodo saltans virus[9]1,385,8691227 proteins (predicted)~300yes (~40 nm)MF782455
Megavirus chilensis[10]1,259,1971120 proteins (predicted)440yes (75 nm)JN258408
Mamavirus[11]1,191,6931023 proteins (predicted)500yes (120 nm)JF801956
Mimivirus[12][13]1,181,549979 proteins 39 non-coding500yes (120 nm)NC_014649
Tupanvirus[14]1,500,0001276-1425 proteins≥450+550[15]KY523104
MF405918[16]

The whole list is in the Giant Virus Toplist created by the Giant Virus Finder software.[17]

Table 2 - Specific common features among giant viruses
Giant virus nameAminoacyl-tRNA synthetaseOctocoral-like 1MutS2Stargate[18]Known virophage[19]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)

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.

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See also

References
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  15. head and tail, respectively
  16. soda lake and deep ocean species of Tupanvirues, respectively
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  19. Fischer MG, Suttle CA (April 2011). "A virophage at the origin of large DNA transposons". Science. 332 (6026): 231–4. Bibcode:2011Sci...332..231F. doi:10.1126/science.1199412. PMID 21385722.
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