ンブスウイルス科

関: dianthovirus、tomato bushy stunt virus、tombusvirus

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2015/12/16 08:47:59」(JST)

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Tombusviridae is a family of single-stranded positive sense RNA plant viruses. There are currently 71 species in this family, divided among 13 genera.^[1]^[2] The name is derived from the type species of the Tombusvirus genus, Tomato bushy stunt virus (TBSV).^[3]

Genome

All Tombusviridae have a non-segmented linear genome, with the exception of Dianthoviruses, whose genome is bipartite.^[4] The genome is approximately 4.6-4.8kb in length, with a 5' cap, and it encodes 4-6 ORFs. The polymerase encodes an amber stop codon which is the site of a readthrough event within ORF1, producing two products necessary for replication. There is no helicase encoded by the virus.

Structure

The RNA is encapsulated in an icosahedral (T=3) capsid, composed of 180 units of a single coat protein 27–42K in size; the virion measures 28–35 nm in diameter, and it is not enveloped.^[1]^[5]

Genus	Structure	Symmetry	Capsid	Genomic Arrangement	Genomic Segmentation
Tombusvirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Gallantivirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Macanavirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Dianthovirus	Icosahedral	T=3	Non-Enveloped	Linear	Bipartite
Carmovirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Alphanecrovirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Avenavirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Panicovirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Betanecrovirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Aureusvirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Umbravirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Machlomovirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite
Zeavirus	Icosahedral	T=3	Non-Enveloped	Linear	Monopartite

Life Cycle

Viral replication is cytoplasmic, and is lysogenic. Entry into the host cell is achieved by penetration into the host cell. Replication follows the positive stranded RNA virus replication model. Positive stranded RNA virus transcription, using the premature termination model of subgenomic RNA transcription is the method of transcription. Translation takes place by leaky scanning, -1 ribosomal frameshifting, viral initiation, and suppression of termination. The virus exits the host cell by tubule-guided viral movement. Plants serve as the natural host. Transmission routes are mechanical, seed borne, and contact.^[1]

Viruses in this family are primarily soil-borne, some transmitted by fungal species of the order Chytridiales, others by no known vector. Virions may spread by water, root growth into infected soil, contact between plants, pollen, or seed, depending on the virus species. These viruses may be successfully transmitted by grafting or mechanical inoculation, and both the virion and the genetic material alone are infective.^[5]

Genus	Host Details	Tissue Tropism	Entry Details	Release Details	Replication Site	Assembly Site	Transmission
Tombusvirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Gallantivirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Macanavirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Dianthovirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Carmovirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Alphanecrovirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Avenavirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Panicovirus	Plants: panicae	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Betanecrovirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Aureusvirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Umbravirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Machlomovirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed
Zeavirus	Plants	None	Viral movement; mechanical innoculation	Viral movement	Cytoplasm	Cytoplasm	Mechanical: contact; seed

Replication

Members of Tombusviridae replicate in the cytoplasm, by use of negative strand templates. The replication process leaves a surplus of positive sense (+)RNA strands, and it is thought that not only does the viral RNA act as a template for replication, but is also able to manipulate and regulate RNA synthesis.^{[citation needed]}

The level of RNA synthesis has been shown to be affected by the cis-acting properties of certain elements on the RNA (such as RNA1 and 2^[6]^[7]), which include core promoter sequences which regulate the site of initiation for the complementary RNA strand synthesis. This mechanism is thought to be recognised by RNA-dependent RNA polymerase, found encoded within the genome.^{[citation needed]}

Tombusviridae have been found to co-opt GAPDH, a host metabolic enzyme, for use in the replication center. GAPDH may bind to the (-)RNA strand and keep it in the replicase complex, allowing (+)RNA strands synthesized from it to be exported and accumulate in the host cell. Downregulation of GAPDH reduced viral RNA accumulation, and eliminated the surplus of (+)RNA copies.^[8]

Notes

Research has shown that infection of plants from tombusviruses contain defective interfering RNAs that are born directly from the viruses RNA genome, and no host genome. Viral DI RNAs with their small size and cis-acting elements are good templates both in vivo and in vitro on which to study RNA replication.^{[citation needed]}

Sub-genomic RNA is used in the synthesis of some proteins; they are generated by premature termination of (-)strand synthesis. sgRNAs and sgRNA negative-sense templates are found in infected cells.^[5]

Taxonomy

Group: ssRNA(+)

Order: Unassigned

Family: Tombusviridae
- Genus: Alphanecrovirus
  Olive latent virus 1
  
  Olive mild mosaic virus
  
  Tobacco necrosis virus A
- Genus: Aureusvirus
  Cucumber leaf spot virus
  
  Johnsongrass chlorotic stripe mosaic virus
  
  Maize white line mosaic virus
  
  Pothos latent virus
- Genus: Avenavirus
  Oat chlorotic stunt virus
- Genus: Betanecrovirus
  Beet black scorch virus
  
  Leek white stripe virus
  
  Tobacco necrosis virus D
- Genus: Carmovirus
  Ahlum waterborne virus
  
  Angelonia flower break virus
  
  Bean mild mosaic virus
  
  Calibrachoa mottle virus
  
  Cardamine chlorotic fleck virus
  
  Carnation mottle virus
  
  Cowpea mottle virus
  
  Cucumber soil-borne virus
  
  Hibiscus chlorotic ringspot virus
  
  Honeysuckle ringspot virus
  
  Japanese iris necrotic ring virus
  
  Melon necrotic spot virus
  
  Nootka lupine vein clearing virus
  
  Pea stem necrosis virus
  
  Pelargonium flower break virus
  
  Saguaro cactus virus
  
  Soybean yellow mottle mosaic virus
  
  Turnip crinkle virus
  
  Weddel waterborne virus
- Genus: Dianthovirus
  Carnation ringspot virus
  
  Red clover necrotic mosaic virus
  
  Sweet clover necrotic mosaic virus
- Genus: Gallantivirus
  Galinsoga mosaic virus
- Genus: Macanavirus
  Furcraea necrotic streak virus
- Genus: Machlomovirus
  Maize chlorotic mottle virus
- Genus: Panicovirus
  Cocksfoot mild mosaic virus
  
  Panicum mosaic virus
  
  Thin paspalum asymptomatic virus
- Genus: Tombusvirus
  Artichoke mottled crinkle virus
  
  Carnation Italian ringspot virus
  
  Cucumber Bulgarian virus
  
  Cucumber necrosis virus
  
  Cymbidium ringspot virus
  
  Eggplant mottled crinkle virus
  
  Grapevine Algerian latent virus
  
  Havel River virus
  
  Lato River virus
  
  Limonium flower distortion virus
  
  Moroccan pepper virus
  
  Neckar River virus
  
  Pelargonium leaf curl virus
  
  Pelargonium necrotic spot virus
  
  Petunia asteroid mosaic virus
  
  Sitke waterborne virus
  
  Tomato bushy stunt virus
- Genus: Umbravirus
  Carrot mottle mimic virus
  
  Carrot mottle virus
  
  Groundnut rosette virus
  
  Lettuce speckles mottle virus
  
  Pea enation mosaic virus 2
  
  Tobacco bushy top virus
  
  Tobacco mottle virus
- Genus: Unassigned
  Chenopodium necrosis virus
  
  Elderberry latent virus
  
  Pelargonium chlorotic ring pattern virus
  
  Pelargonium line pattern virus
  
  Pelargonium ringspot virus
  
  Rosa rugosa leaf distortion virus
  
  Trailing lespedeza virus 1
- Genus: Zeavirus
  Maize necrotic streak virus

^[2]

Pelarspovirus is an additional genus that has been proposed.^[9]

References

^ ^a ^b ^c "Viral Zone". ExPASy. Retrieved 15 June 2015.
^ ^a ^b ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.
^ Habili, N. and Symons, R. H. (1989). Evolutionary relationship between luteoviruses and other RNA plant viruses based on sequence motifs in their putative RNA polymerases and nucleic acid helicases. Nucleic Acids Research 17:23, 9543–55
^ Wiley InterScience Encyclopedia of Life Sciences: Tombusviridae
^ ^a ^b ^c ICTVdB—The Universal Virus Database, version 3 00.074. Tombusviridae
^ Lommel SA, Weston-Fina M, Xiong Z, Lomonossoff GP (September 1988). "The nucleotide sequence and gene organization of red clover necrotic mosaic virus RNA-2". Nucleic Acids Res. 16 (17): 8587–602. doi:10.1093/nar/16.17.8587. PMC 338578. PMID 3047682.
^ Mizumoto H, Tatsuta M, Kaido M, Mise K, Okuno T (November 2003). "Cap-independent translational enhancement by the 3' untranslated region of red clover necrotic mosaic virus RNA1". J. Virol. 77 (22): 12113–21. doi:10.1128/JVI.77.22.12113-12121.2003. PMC 254280. PMID 14581548.
^ Wang, R. and Nagy, P. (2008) Tomato bushy stunt virus Co-Opts the RNA-Binding Function of a Host Metabolic Enzyme for Viral Genomic RNA Synthesis. Cell Host & Microbe 3:3 178–187
^ Castaño A, Ruiz L, Hernández C (2009) Insights into the translational regulation of biologically active open reading frames of Pelargonium line pattern virus. Virology 386(2):417–426

External links

Viralzone: Tombusviridae
ICTV

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A novel single-stranded RNA virus isolated from the rice-pathogenic fungus Magnaporthe oryzae with similarity to members of the family Tombusviridae.

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Here, we report a novel virus isolated from rice blast fungus, Magnaporthe oryzae, an important plant pathogen. This virus has an RNA genome of 3246 nucleotides. Its genome possesses two in-frame open reading frames (ORFs). The smaller ORF1 encodes a protein with significant similarity to a protein
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A Y-shaped RNA structure in the 3' untranslated region together with the trans-activator and core promoter of Red clover necrotic mosaic virus RNA2 is required for its negative-strand RNA synthesis.

An Mengnan,Iwakawa Hiro-oki,Mine Akira,Kaido Masanori,Mise Kazuyuki,Okuno Tetsuro
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Red clover necrotic mosaic virus (RCNMV) is a positive-strand RNA virus with a bipartite genome. RNA1 encodes N-terminally overlapping replication proteins, p27 and p88. RNA2 is replicated efficiently …
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リンク元	「トンブスウイルス科」「tomato bushy stunt virus」「dianthovirus」「tombusvirus」

「トンブスウイルス科」

Tombusviridae
Virus classification
Group:	Group IV ((+)ssRNA)
Family:	Tombusviridae
Genera
	Betanecrovirus; Aureusvirus; Machlomovirus; Panicovirus; Alphanecrovirus; Umbravirus; Avenavirus; Dianthovirus; Carmovirus; Macanavirus; Zeavirus; Tombusvirus; Gallantivirus;

　　[★]

ラ: Tombusviridae
関: トンブスウイルス、ダイアンソウイルス、トマトブッシースタントウイルス

「tomato bushy stunt virus」

　　[★]

トマトブッシースタントウイルス

関: dianthovirus、Tombusviridae、tombusvirus

「dianthovirus」

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ダイアンソウイルス

関: tomato bushy stunt virus、Tombusviridae、tombusvirus

「tombusvirus」

　　[★]

トンブスウイルス

関: dianthovirus、tomato bushy stunt virus、Tombusviridae

[ViralZone-1] "Viral Zone". ExPASy. Retrieved 15 June 2015.

[ICTV-2] ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.

[3] Habili, N. and Symons, R. H. (1989). Evolutionary relationship between luteoviruses and other RNA plant viruses based on sequence motifs in their putative RNA polymerases and nucleic acid helicases. Nucleic Acids Research 17:23, 9543–55

[wiley-4] Wiley InterScience Encyclopedia of Life Sciences: Tombusviridae

[ictvdb-5] ICTVdB—The Universal Virus Database, version 3 00.074. Tombusviridae

[6] Lommel SA, Weston-Fina M, Xiong Z, Lomonossoff GP (September 1988). "The nucleotide sequence and gene organization of red clover necrotic mosaic virus RNA-2". Nucleic Acids Res. 16 (17): 8587–602. doi:10.1093/nar/16.17.8587. PMC 338578. PMID 3047682.

[7] Mizumoto H, Tatsuta M, Kaido M, Mise K, Okuno T (November 2003). "Cap-independent translational enhancement by the 3' untranslated region of red clover necrotic mosaic virus RNA1". J. Virol. 77 (22): 12113–21. doi:10.1128/JVI.77.22.12113-12121.2003. PMC 254280. PMID 14581548.

[8] Wang, R. and Nagy, P. (2008) Tomato bushy stunt virus Co-Opts the RNA-Binding Function of a Host Metabolic Enzyme for Viral Genomic RNA Synthesis. Cell Host & Microbe 3:3 178–187

[9] Castaño A, Ruiz L, Hernández C (2009) Insights into the translational regulation of biologically active open reading frames of Pelargonium line pattern virus. Virology 386(2):417–426

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Tombusviridae