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Poxviridae

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Poxviruses
Virus classification Edit this classification
Subfamilies

Poxviridae is a family of viruses. Humans, vertebrates, and arthropods serve as natural hosts. There are currently 69 species in this family, divided among 28 genera, which are divided into two subfamilies. Diseases associated with this family include smallpox.[1][2]

Four genera of poxviruses may infect humans: orthopoxvirus, parapoxvirus, yatapoxvirus, molluscipoxvirus. Orthopox: smallpox virus (variola), vaccinia virus, cowpox virus, monkeypox virus; Parapox: orf virus, pseudocowpox, bovine papular stomatitis virus; Yatapox: tanapox virus, yaba monkey tumor virus; Molluscipox: molluscum contagiosum virus (MCV).[3] The most common are vaccinia (seen on Indian subcontinent) and molluscum contagiosum, but monkeypox infections are rising (seen in west and central African rainforest countries). The similarly named disease chickenpox is not a true poxvirus and is actually caused by the herpesvirus varicella zoster.

Phylogeny

A new systematic has been proposed recently after findings of a new squirrel poxvirus in Berlin, Germany.[4]

Structure

A) Electron micrograph of poxvirus particles in synovium of a big brown bat, northwestern United States. B) Negative staining of poxvirus particles in cell culture supernatant. Scale bar = 100 nm.

Poxviridae viral particles (virions) are generally enveloped (external enveloped virion- EEV), though the intracellular mature virion (IMV) form of the virus, which contains different envelope, is also infectious. They vary in their shape depending upon the species but are generally shaped like a brick or as an oval form similar to a rounded brick because they are wrapped by the endoplasmic reticulum. The virion is exceptionally large, its size is around 200 nm in diameter and 300 nm in length and carries its genome in a single, linear, double-stranded segment of DNA.[5] By comparison, Rhinovirus is 1/10 as large as a typical Poxviridae virion.[6]

Genus Structure Symmetry Capsid Genomic arrangement Genomic segmentation
Betaentomopoxvirus Ovoid Enveloped Linear Monopartite
Yatapoxvirus Brick-shaped Enveloped Linear Monopartite
Cervidpoxvirus Brick-shaped Enveloped Linear Monopartite
Gammaentomopoxvirus Ovoid Enveloped Linear Monopartite
Leporipoxvirus Brick-shaped Enveloped Linear Monopartite
Suipoxvirus Brick-shaped Enveloped Linear Monopartite
Molluscipoxvirus Brick-shaped Enveloped Linear Monopartite
Crocodylidpoxvirus Ovoid Enveloped Linear Monopartite
Alphaentomopoxvirus Ovoid Enveloped Linear Monopartite
Capripoxvirus Brick-shaped Enveloped Linear Monopartite
Orthopoxvirus Brick-shaped Enveloped Linear Monopartite
Avipoxvirus Brick-shaped Enveloped Linear Monopartite
Parapoxvirus Ovoid Enveloped Linear Monopartite

Replication

Replication of the poxvirus involves several stages. The virus first binds to a receptor on the host cell surface; the receptors for the poxvirus are thought to be glycosaminoglycans (GAGs). After binding to the receptor, the virus enters the cell where it uncoats. Uncoating of the virus is a two step process. Firstly the outer membrane is removed as the particle enters the cell; secondly the virus particle (without the outer membrane) fuses with the cellular membrane to release the core into the cytoplasm. The pox viral genes are expressed in two phases. The early genes encode the non-structural protein, including proteins necessary for replication of the viral genome, and are expressed before the genome is replicated. The late genes are expressed after the genome has been replicated and encode the structural proteins to make the virus particle. The assembly of the virus particle occurs in five stages of maturation that lead to the final exocytosis of the new enveloped virion. After the genome has been replicated, the immature virion (IV) assembles the A5 protein to create the intracellular mature virion (IMV). The protein aligns and the brick-shaped envelope of the intracellular enveloped virion (IEV). These IEV particles are then fused to the cell plasma to form the cell-associated enveloped virion (CEV). Finally, the CEV encounters the microtubules and the virion prepares to exit the cell as an extracellular enveloped virion (EEV). The assembly of the virus particle occurs in the cytoplasm of the cell and is a complex process that is currently being researched to understand each stage in more depth. Considering the fact that this virus is large and complex, replication is relatively quick taking approximately 12 hours until the host cell dies by the release of viruses.

The replication of poxvirus is unusual for a virus with double-stranded DNA genome (dsDNA) because it occurs in the cytoplasm,[7] although this is typical of other large DNA viruses.[8] Poxvirus encodes its own machinery for genome transcription, a DNA dependent RNA polymerase,[9] which makes replication in the cytoplasm possible. Most dsDNA viruses require the host cell's DNA-dependent RNA polymerase to perform transcription. These host DNA are found in the nucleus, and therefore most dsDNA viruses carry out a part of their infection cycle within the host cell's nucleus.

Genus Host details Tissue tropism Entry details Release details Replication site Assembly site Transmission
Betaentomopoxvirus Lepidoptera insects; orthoptera insects Hemocytes; fat bodies Glycosaminoglycans Lysis; budding; occlusion Cytoplasm Cytoplasm Unknown
Yatapoxvirus Monkeys; baboons None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Contact; insects
Cervidpoxvirus Deer None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Unknown
Gammaentomopoxvirus Insects Hemocytes; fat bodies Glycosaminoglycans Lysis; budding; occlusion Cytoplasm Cytoplasm Unknown
Leporipoxvirus Lagomorph; squirrels None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Arthropods; contact
Suipoxvirus Swine None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Unknown
Molluscipoxvirus Humans None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Sex; contact
Crocodylidpoxvirus Crocodiles None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Unknown
Alphaentomopoxvirus Monkeys; baboons None Glycosaminoglycans Lysis; budding; occlusion Cytoplasm Cytoplasm Contact; insects
Capripoxvirus Sheep; goat; cattle None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Arthropods; contact
Orthopoxvirus Humans; mammals None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Variola virus; Respiratory; contact; zoonosis
Avipoxvirus Birds None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Arthropods; aerosol
Parapoxvirus Humans; mammals None Glycosaminoglycans Lysis; budding Cytoplasm Cytoplasm Zoonosis; contact

Evolution

The ancestor of the poxviruses is not known but structural studies suggest it may have been an adenovirus or a species related to both the poxviruses and the adenoviruses.[10]

Based on the genome organisation and DNA replication mechanism it seems that phylogenetic relationships may exist between the rudiviruses (Rudiviridae) and the large eukaryal DNA viruses: the African swine fever virus (Asfarviridae), Chlorella viruses (Phycodnaviridae) and poxviruses (Poxviridae).[11]

The mutation rate in these genomes has been estimated to be 0.9-1.2 x 10−6 substitutions per site per year.[12] A second estimate puts this rate at 0.5-7 × 10−6 nucleotide substitutions per site per year.[13] A third estimate places the rate at 4-6 × 10−6.[14]

The last common ancestor of the extant poxviruses that infect vertebrates existed 0.5 million years ago. The genus Avipoxvirus diverged from the ancestor 249 ± 69 thousand years ago. The ancestor of the genus Orthopoxvirus was next to diverge from the other clades at 0.3 million years ago. A second estimate of this divergence time places this event at 166,000 ± 43,000 years ago.[13] The division of the Orthopox into the extant genera occurred ~14,000 years ago. The genus Leporipoxvirus diverged ~137,000 ± 35,000 years ago. This was followed by the ancestor of the genus Yatapoxvirus. The last common ancestor of the Capripoxvirus and Suipoxvirus diverged 111,000 ± 29,000 years ago.

An isolate from a fish - Salmon Gill Poxvirus - appears to be the earliest branch in the Chordopoxvirinae.[15]

Smallpox

The date of the appearance of smallpox is not settled. It most likely evolved from a rodent virus between 68,000 and 16,000 years ago.[16][17] The wide range of dates is due to the different records used to calibrate the molecular clock. One clade was the variola major strains (the more clinically severe form of smallpox) which spread from Asia between 400 and 1,600 years ago. A second clade included both alastrim minor (a phenotypically mild smallpox) described from the American continents and isolates from West Africa which diverged from an ancestral strain between 1,400 and 6,300 years before present. This clade further diverged into two subclades at least 800 years ago.

A second estimate has placed the separation of variola from Taterapox at 3000–4000 years ago.[14] This is consistent with archaeological and historical evidence regarding the appearance of smallpox as a human disease which suggests a relatively recent origin. However, if the mutation rate is assumed to be similar to that of the herpesviruses the divergence date between variola from Taterapox has been estimated to be 50,000 years ago.[14] While this is consistent with the other published estimates it suggests that the archaeological and historical evidence is very incomplete. Better estimates of mutation rates in these viruses are needed.

Taxonomy

Group: dsDNA

Order: Unassigned

[2]

The name of the family, Poxviridae, is a legacy of the original grouping of viruses associated with diseases that produced poxes in the skin. Modern viral classification is based on phenotypic characteristics; morphology, nucleic acid type, mode of replication, host organisms, and the type of disease they cause. The smallpox virus remains as the most notable member of the family.

The species in the subfamily Chordopoxvirinae infect vertebrates and those in the subfamily Entomopoxvirinae infect insects. There are 10 recognised genera in the Chordopoxvirinae and 3 in the Entomopoxvirinae. Both subfamiles also contain a number of unclassified species for which new genera may be created in the future. Cotia virus is an unusual virus that may belong to a new genus.[18] Two more poxviruses are NY_014 and Murmansk poxvirus.[19]

Notes

The GC-content of these genomes differs considerably.[20] Avipoxvirus, Capripoxvirus, Cervidpoxvirus, Orthopoxvirus, Suipoxvirus, Yatapoxvirus and one Entomopox genus (Betaentomopoxvirus) along with several other unclassified Entomopoxviruses have a low G+C content while others - Molluscipoxvirus, Orthopoxvirus, Parapoxvirus and some unclassified Chordopoxvirus - have a relatively high G+C content. The reasons for these differences are not known.

Phylogenetic analysis of 26 Chordopoxviruses genomes has shown that the central region of the genome is conserved and contains ~90 genes.[21] The termini in contrast are not conserved between species. Of this group Avipoxvirus is the most divergent. The next most divergent is Molluscipoxvirus. Capripoxvirus, Leporipoxvirus, Suipoxvirus and Yatapoxvirus genera cluster together: Capripoxvirus and Suipoxvirus share a common ancestor and are distinct from the genus Orthopoxvirus. Within the Othopoxvirus genus Cowpox virus strain Brighton Red, Ectromelia virus and Monkeypox virus do not group closely with any other member. Variola virus and Camelpox virus form a subgroup. Vaccinia virus is most closely related to CPV-GRI-90.

Vaccinia virus

Main article: Vaccinia virus

The prototypial poxvirus is vaccinia virus, known for its role as the active agent in the eradication of smallpox. The vaccinia virus is an effective tool for foreign protein expression, as it elicits a strong host immune-response. The vaccinia virus enters cells primarily by cell fusion, although currently the receptor responsible is unknown.

Vaccinia contains three classes of genes: early, intermediate and late. These genes are transcribed by viral RNA polymerase and associated transcription factors. Vaccinia replicates its genome in the cytoplasm of infected cells, and after late-stage gene expression undergoes virion morphogenesis, which produces IMV contained within an envelope membrane. The exact origin of the envelope membrane is still unknown. The IMV is then transported to the Golgi apparatus where it is wrapped with an additional two membranes, becoming the Intracellular Enveloped Virus (IEV). The IEV is transported along cytoskeletal microtubules to reach the cell periphery, where it fuses with the plasma membrane to become the Cell-associated Enveloped Virus (CEV). This triggers actin tails on cell surfaces or is released as EEV.

History

Diseases caused by pox viruses, especially smallpox, have been known about for centuries. One of the earliest suspected cases is that of Egyptian pharaoh Ramses V who is thought to have died from smallpox circa 1150 years BCE.[22][23] Smallpox was thought to have been transferred to Europe around the early 8th century and then to the Americas in the early 16th century. It is widely accepted that the main defeat of the Aztecs was due to a smallpox epidemic and within two years over 3.2 million Aztecs died. This death toll can be attributed to the American population's complete lack of exposure to the virus over millennia. A century after Edward Jenner showed that the less potent cow pox could be used to effectively vaccinate against the more deadly smallpox, a worldwide effort to vaccinate everyone against smallpox began with the ultimate goal to rid the world of the plague-like epidemic. The last case of endemic smallpox occurred in Somalia in 1977. Extensive searches over two years detected no further cases, and in 1979 the World Health Organization (WHO) declared the disease officially eradicated. In 1986, all virus samples were destroyed or transferred to two approved WHO reference labs: at the headquarters of the federal Centers for Disease Control and Prevention (the C.D.C.) in Atlanta, Georgia (the United States) and at the Institute of Virus Preparations in Moscow.[24] Post September 11, 2001 the American and UK governments have had increased concern over the use of smallpox, or a smallpox like disease, in bio-terrorism.

References

  1. ^ "Viral Zone". ExPASy. Retrieved 15 June 2015.
  2. ^ a b ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.
  3. ^ "Pathogenic Molluscum Contagiosum Virus Sequenced". Antiviral Agents Bulletin: 196–7. August 1996. Retrieved 2006-07-16.
  4. ^ Wibbelt, Gudrun; Tausch, Simon H.; Dabrowski, Piotr W.; Kershaw, Olivia; Nitsche, Andreas; Schrick, Livia (2017). "Berlin Squirrelpox Virus, a New Poxvirus in Red Squirrels, Berlin, Germany". Emerging Infectious Diseases. 23 (10): 1726–1729. doi:10.3201/eid2310.171008. PMC 5621524. PMID 28930029.(for systematic see figure 2)
  5. ^ International Committee on Taxonomy of Viruses (2004-06-15). "ICTVdb Descriptions: 58. Poxviridae". Retrieved 2005-02-26.
  6. ^ How Big is a ... ? at Cells Alive!. Retrieved 2005-02-26.
  7. ^ Mutsafi, Y; Zauberman, N; Sabanay, I; Minsky, A (30 Mar 2010). "Vaccinia-like cytoplasmic replication of the giant Mimivirus". Proceedings of the National Academy of Sciences USA. 107 (13): 5978–82. doi:10.1073/pnas.0912737107. PMC 2851855. PMID 20231474..
  8. ^ Racaniello, Vincent (4 March 2014). "Pithovirus: Bigger than Pandoravirus with a smaller genome". Virology Blog. Retrieved 4 March 2014.
  9. ^ National Center for Infectious Diseases, Centers for Disease Control and Prevention, 1600 Clifton Rd., Atlanta, GA 30333, USA. "DNA-dependent RNA polymerase rpo35 (Vaccinia virus)". National Center for Biotechnology Information (NCBI), NIH, Bethesda, MD, USA.{{cite web}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  10. ^ Bahar, MW; Graham, SC; Stuart, DI; Grimes, JM (2011). "Insights into the evolution of a complex virus from the crystal structure of vaccinia virus D13". Structure. 19 (7): 1011–1020. doi:10.1016/j.str.2011.03.023. PMC 3136756. PMID 21742267.
  11. ^ Prangishvili, D; Garrett, RA (2004). "Exceptionally diverse morphotypes and genomes of crenarchaeal hyperthermophilic viruses". Biochem Soc Trans. 32 (2): 204–208. doi:10.1042/bst0320204. PMID 15046572.
  12. ^ Babkin IV, Shchelkunov SN (2006) The time scale in poxvirus evolution. Mol Biol (Mosk) 40(1):20-24
  13. ^ a b Babkin, IV; Babkina, IN (2011). "Molecular dating in the evolution of vertebrate poxviruses". Intervirology. 54 (5): 253–260. doi:10.1159/000320964. PMID 21228539.
  14. ^ a b c Hughes, AL; Irausquin, S; Friedman, R (2010). "The evolutionary biology of poxviruses". Infect Genet Evol. 10 (1): 50–59. doi:10.1016/j.meegid.2009.10.001. PMC 2818276. PMID 19833230.
  15. ^ Gjessing, MC; Yutin, N; Tengs, T; Senkevich, T; Koonin, E; Rønning, HP; Alarcon, M; Ylving, S; Lie, KI; Saure, B; Tran, L; Moss, B; Dale, OB (2015). "Salmon Gill Poxvirus, the Deepest Representative of the Chordopoxvirinae". J Virol. 89 (18): 9348–9867. doi:10.1128/JVI.01174-15. PMC 4542343. PMID 26136578.
  16. ^ Esposito, JJ; Sammons, SA; Frace, AM; Osborne, JD; Olsen-Rasmussen, M; Zhang, M; Govil, D; Damon, IK; et al. (August 2006). "Genome sequence diversity and clues to the evolution of variola (smallpox) virus". Science (Submitted manuscript). 313 (5788): 807–812. doi:10.1126/science.1125134. PMID 16873609. {{cite journal}}: Explicit use of et al. in: |first9= (help)
  17. ^ Li, Y; Carroll, DS; Gardner, SN; Walsh, MC; Vitalis, EA; Damon, IK (2007). "On the origin of smallpox: correlating variola phylogenics with historical smallpox records". Proc Natl Acad Sci USA. 104 (40): 15787–15792. doi:10.1073/pnas.0609268104. PMC 2000395. PMID 17901212.
  18. ^ Afonso PP, Silva PM, Schnellrath LC, Jesus DM, Hu J, Yang Y, Renne R, Attias M, Condit RC, Moussatché N, Damaso CR (2012) Biological characterization and next-generation genome sequencing of the unclassified Cotia virus SPAn232 (Poxviridae). J Virol
  19. ^ Smithson C, Meyer H, Gigante CM, Gao J, Zhao H, Batra D, Damon I, Upton C, Li Y (2017) Two novel poxviruses with unusual genome rearrangements: NY_014 and Murmansk. Virus Genes
  20. ^ Roychoudhury, S; Pan, A; Mukherjee, D (2011). "Genus specific evolution of codon usage and nucleotide compositional traits of poxviruses". Virus Genes. 42 (2): 189–199. doi:10.1007/s11262-010-0568-2. PMID 21369827.
  21. ^ Gubser, C; Hué, S; Kellam, P; Smith, GL (2004). "Poxvirus genomes: a phylogenetic analysis". J Gen Virol. 85 (1): 105–117. doi:10.1099/vir.0.19565-0. PMID 14718625.
  22. ^ Hopkins, Donald R. (2002) [1983]. The greatest killer: smallpox in history, with a new introduction. University of Chicago Press. p. 15. By special permission of the late President Anwar el Sadat, I was allowed to examine the front upper half of Ramses V's unwrapped mummy in the Cairo Museum in 1979. …Inspection of the mummy revealed a rash of elevated "pustules," each about two to four millimeters in diameter, …(An attempt to prove that this rash was caused by smallpox by electron-microscopic examination of tiny pieces of tissue that had fallen on the shroud was unsuccessful. I was not permitted to excise one of the postules.) …The appearance of the larger pustules and the apparent distribution of the rash are similar to smallpox rashes I have seen in more recent victims
  23. ^ Date of Ramses V's death derived from the Encyclopedia of Ancient Egypt, Margaret Bunson (New York: Facts On File, 2002) ISBN 0816045631 p.337.
  24. ^ Henderson, D. A.; Inglesby, Thomas V.; Bartlett, John G.; Ascher, Michael S.; Eitzen, Edward; Jahrling, Peter B.; Hauer, Jerome; Layton, Marcelle; McDade, Joseph; Osterholm, Michael T.; O'Toole, Tara; Parker, Gerald; Perl, Trish; Russell, Philip K.; Tonat, Kevin; For The Working Group On Civilian Biodefense (1999). "Smallpox as a Biological Weapon: Medical and Public Health Management". JAMA: The Journal of the American Medical Association. 281 (22): 2127–37. doi:10.1001/jama.281.22.2127. PMID 10367824.
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