Ordine Mononegavirales Classificazione

Gruppo: III (dsRNA)

Ordine:Mononegavirales

Mononegavirales è un ordine di virus con genoma a RNA a singolo filamento negativo, suddiviso

in quattro famiglie comprendenti un ampio numero di agenti di svariate malattie infettive degli

animali, uomo compreso, e delle piante.

Al microscopio elettronico i virioni appaiono costituiti da un nucleocapside a simmetria elicoidale

circondato da un rivestimento lipidico, sensibile all’etere, in cui sono inserite glicoproteine virali.

Di dimensioni e forma varia (filamentosi, o a forma di proiettile, o sferici o pleomorfi). L’RNA

virale (un’unica molecola, non segmentata) costituisce solo lo 0,5% della particella infettiva. Il

genoma virale codifica per proteine strutturali e per proteine non strutturali. Il 15-25% in peso dei

virioni è costituito da lipidi.

Le famiglie appartenenti a quest’ordine sono:

• Bornaviridae

• Filoviridae

• Paramyxoviridae

• Rhabdoviridae

Nella tabella 1 sono riportate, in sintesi, le caratteristiche delle quattro famiglie.

Tabella 1 – Sintesi delle caratteristiche delle quattro famiglie dell’ordine Mononegavirales.

Le famiglie, sottofamiglie, generi e epecie dell’ordine Mononegavirales sono di seguito riportati.

Famiglia Bornaviridae: è una famiglia di virus neurotropi, in possesso di un breve genoma ad RNA

a singolo filamento negativo, in grado di replicarsi all’interno del nucleo della cellula ospite.

Genere Bornavirus: in possesso di un breve genoma ad RNA a singolo filamento negativo in

grado di replicarsi all’interno del nucleo della cellula ospite. Vi appartengono l’agente

eziologico della malattia di Borna, una malattia infettiva mortale negli animali

(soprattutto cavallo e pecora, ma anche cani, gatti e bovini) e gli agenti eziologici della

dilatazione proventricolare negli uccelli.

Specie Borna disease virus *

Famiglia Filoviridae: è una famiglia di virus RNA che comprende tre generi di virus

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Genere Cuevavirus

Specie Cuevavirus Lloviu

Genere Marburgvirus

Specie Lake Victoria virus *

Genere Ebolavirus

Specie Zaire virus *

Specie Bundibugyo virus

Specie Ivory Coast virus

Specie Reston virus

Specie Taï Forest virus

Specie Sudan virus

Famiglia Paramyxoviridae: è una famiglia di virus pleomorfi in possesso di un genoma ad RNA a

singolo filamento negativo avvolto da una membrana lipoproteica. Appartengono a questa

famiglia, fra gli altri, i virus parainfluenzali, il virus della parotite, il virus del morbillo e il virus

respiratorio sinciziale umano. Possiede due sottofamiglie.

Sottofamiglia Paramyxovirinae

Genere Avulavirus

Genere Henipavirus

Genere Morbillivirus

Genere Respirovirus

Genere Rubulavirus

Sottofamiglia Pneumovirinae:

Genere Pneumovirus

Genere Metapneumovirus

Altri virus della famiglia Paramyxoviridae non assegnati ad alcun genere sono:

• Tupaia paramyxovirus

• Fer-de-Lance virus

• Menangle virus

• Nariva virus

• Tioman virus

Famiglia Rhabdoviridae: è costituita da virus altamente infettanti, agenti di malattie degli animali e

delle piante, generalmente trasmessi da artropodi. Sono tra i virus più diffusi in natura; oltre ai

virus delle piante, sono stati riconosciuti oltre 70 rhabdovirus dei vertebrati, per non dire delle

centinaia che aspettano ancora di essere identificati. Si tratta di particelle di forma allungata

della lunghezza variabile da 100 a 430 nm e larghezza intorno a 60-85 nm. I virus degli animali

hanno una caratteristica forma "a proiettile", con una estremità piatta e l'altra arrotondata. I virus

delle piante sono invece bastoncellari, con le due estremità arrotondate. Sono virus provvisti di

capside, con particolari estroflessioni a punta. All’interno è contenuto il ribonucleocapside dal

caratteristico aspetto striato. L’acido nucleico è costituito da RNA monocatenario, a polarità

negativa. Il più importante e noto membro della famiglia è il virus della rabbia.

A questa famiglia appartengono i seguenti generi di virus:

• Genere Lyssavirus (figura 1A), con i seguenti virus:

1. Aravan virus (ARAV)

2. Australian bat lyssavirus (ABLV)

3. Bokeloh bat virus (BBLV)

4. Duvenhage virus (DUVV)

5. European bat virus 1 (EBLV-1)

6. European bat virus 2 (EBLV-2)

7. Ikoma virus

8. Irkut virus

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9. Khujand virus (KHUV)

10. Lagos bat virus (LBV)

11. Mokola virus (MOKV)

12. Rabies virus (RABV)*

13. Shimoni bat virus (SHBV)

14. West Caucasian bat virus (WCBV)

• Genere Cytorhabdovirus (figura 1B), con i seguenti virus:

1. Alfalfa dwarf virus (ADV)

2. Barley yellow striate mosaic virus (BYSMV)

3. Broccoli necrotic yellows virus (BNYV)

4. Colocasia bobone disease-associated virus (CBDaV)

5. Festuca leaf streak virus (FLSV)

6. Lettuce necrotic yellows virus (LNYV)*

7. Lettuce yellow mottle virus (LYMoV)

8. Northern cereal mosaic virus (NCMV)

9. Sonchus virus (SonV)

10. Strawberry crinkle virus (SCV)

11. Wheat American striate mosaic virus (WASMV)

• Genere Ephemerovirus (figura 1C), con i seguenti virus;

1. Adelaide River virus

2. Berrimah virus

3. Bovine ephemeral fever virus *

• Genere Novirhabdovirus (figura 1D), con i seguenti virus:

1. Hirame rhabdovirus (HIRV),

2. Oncorhynchus 1 novirhabdovirus* Infectious hematopoietic necrosis virus (IHNV),

3. Oncorhynchus 2 novirhabdovirus Viral hemorrhagic septicemia virus (VHSV),

4. Snakehead virus (SHRV).

• Genere Nucleorhabdovirus (figura 1E), con i seguenti virus:

1. Datura yellow vein virus

2. Eggplant mottled dwarf virus

3. Maize fine streak virus

4. Maize Iranian mosaic virus

5. Maize mosaic virus

6. Potato yellow dwarf virus *

7. Rice yellow stunt virus

8. Sonchus yellow net virus

9. Sowthistle yellow vein virus

10. Taro vein chlorosis virus

• Genere Vesiculovirus (figura 1F), con i seguenti virus:

1. Carajas virus

2. Chandipura virus

3. Cocal virus

4. Isfahan virus

5. Maraba virus

6. Piry virus

7. Vesicular stomatitis Alagoas virus

8. Vesicular stomatitis Indiana virus *

9. Vesicular stomatitis New Jersey virus

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Seguono le immagini di alcune specie virali della famiglia Rhabdoviridae.

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Figura 1 - Generi Lyssavirus (A), Cytorhabdovirus (B), Ephemerovirus (C), Novirhabdovirus (D),

Nucleorhabdovirus (E) e Vesiculovirus (F).

Bibliografia

1. Adams M.J, Christian P, Ghabrial SA, Knowles NJ, Lavigne R, 2011. The unassigned

viruses, 1197–1207. In King AMQ, Lefkowitz E, Adams MJ, Carstens AMQ, Adams MJ,

Carstens EB, editors. (ed), Virus taxonomy: ninth report of the International Committee on

Taxonomy of Viruses. Elsevier, San Diego, CA.

2. Badrane H, Tordo N, 2001. Host switching in Lyssavirus history from the Chiroptera to the

Carnivora orders. J. Virol. 75, 8096-8104.

3. Balachandran S, Barber GN, 2000. Vesicular stomatitis virus (VSV) therapy of tumors.

IUBMB Life 50, 135-138.

4. Banyard AC, Hayman D, Johnson N, McElhinney L, Fooks AR, 2011.: Bats and

lyssaviruses. Adv. Virus Res. 79, 239-289.

5. Basak S, Mondal A, Polley S, Mukhopadhyay S, Chattopadhyay D, 2007. Reviewing

Chandipura: a vesiculovirus in human epidemics. Biosci. Rep. 27, 275-298.

6. Blondel D, Regad T, Poisson N, Pavie B, Harper F, Pandolfi PP, De The H, Chelbi-Alix MK,

2002. Rabies virus P and small P products interact directly with PML and reorganize PML

nuclear bodies. Oncogene 21, 7957-7970.

7. Bloor S, Maelfait J, Krumbach R, Beyaert R, Randow F, 2010. Endoplasmic reticulum

chaperone gp96 is essential for infection with vesicular stomatitis virus. Proc. Natl. Acad. Sci.

U. S. A. 107, 6970-6975.

8. Botvinkin AD, Poleschuk EM, Kuzmin IV, Borisova TI, Gazaryan SV, Yager P, Rupprecht

CE, 2003. Novel lyssaviruses isolated from bats in Russia. Emerg. Infect. Dis. 9, 1623-1625.

9. Brown DA, Crise B, Rose JK, 1989. Mechanism of membrane anchoring affects polarized

expression of two proteins in MDCK cells. Science 245, 1499-1501.

10. Brzozka K, Finke S, Conzelmann KK, 2005. Identification of the rabies virus alpha/beta

interferon antagonist: phosphoprotein P interferes with phosphorylation of interferon

regulatory factor 3. J. Virol. 79, 7673-7681.

6

11. Brzozka K, Finke S, Conzelmann KK, 2006. Inhibition of interferon signaling by rabies virus

phosphoprotein P: activation-dependent binding of STAT1 and STAT2. J. Virol. 80, 2675-

2683.

12. Chen Z, Green TJ, Luo M, Li H, 2004. Visualizing the RNA molecule in the bacterially

expressed vesicular stomatitis virus nucleoprotein-RNA complex. Structure 12, 227-235.

13. Chenik M, Schnell M, Conzelmann KK, Blondel D, 1998. Mapping the interacting domains

between the rabies virus polymerase and phosphoprotein. J. Virol. 72, 1925-1930.

14. Cleaveland S, Fevre EM, Kaare M, Coleman PG, 2002. Estimating human rabies mortality in

the United Republic of Tanzania from dog bite injuries. Bull. World Health Organ. 80, 304-310.

15. Colonno RJ, Banerjee AK, 1978. Complete nucleotide sequence of the leader RNA synthesized

in vitro by vesicular stomatitis virus. Cell 15, 93-101.

16. Comer JA, Tesh RB, 1991. Phlebotomine sand flies as vectors of vesiculoviruses: a review.

Parassitologia 33 Suppl, 143-150.

17. Connor JH, McKenzie MO, Lyles DS, 2006. Role of residues 121 to 124 of vesicular

stomatitis virus matrix protein in virus assembly and virus-host interaction. J. Virol. 80, 3701-

3711.

18. Conzelmann KK, Schnell M, 1994. Rescue of synthetic genomic RNA analogs of rabies virus

by plasmid-encoded proteins. J. Virol. 68, 713-719.

19. Craven RC, Harty RN, Paragas J, Palese P, Wills JW, 1999. Late domain function identified

in the vesicular stomatitis virus M protein by use of rhabdovirus-retrovirus chimeras. J. Virol.

73, 3359-3365.

20. Da Poian AT, Carneiro FA, Stauffer F, 2005. Viral membrane fusion: is glycoprotein G of

rhabdoviruses a representative of a new class of viral fusion proteins? Braz. J. Med. Biol. Res.

38, 813-823.

21. Desforges M, Despars G, Berard S, Gosselin M, McKenzie MO, Lyles DS, Talbot PJ,

Poliquin L, 2002. Matrix protein mutations contribute to inefficient induction of apoptosis

leading to persistent infection of human neural cells by vesicular stomatitis virus. Virology

295, 63-73.

22. Dietzgen RG, Calisher CH, Kurath G, Kuzmin IV, Rodriguez LL, Stone DM, Tesh RB, Tordo

N, Walker PJ, Wetzel T, Whitfield AE, 2011. Family Rhabdoviridae, King AMQ, Adams MJ,

Carstens EB, Lefkowitz EJ (eds), Virus Taxonomy - 9th Report of the ICTV. Elsevier

Academic Press.

23. Dietzschold B, Wunner WH, Wiktor TJ, Lopes AD, Lafon M, Smith CL, Koprowski H, 1983.

Characterization of an antigenic determinant of the glycoprotein that correlates with

pathogenicity of rabies virus. Proc. Natl. Acad. Sci. U. S. A. 80, 70-74.

24. Dotti CG, Simons K, 1990. Polarized sorting of viral glycoproteins to the axon and dendrites

of hippocampal neurons in culture. Cell 62, 63-72.

25. Durrer P, Gaudin Y, Ruigrok RW, Graf R, Brunner J, 1995. Photolabeling identifies a

putative fusion domain in the envelope glycoprotein of rabies and vesicular stomatitis viruses.

J. Biol. Chem. 270, 17575-17581.

26. Emerson SU, Schubert M, 1987.: Location of the binding domains for the RNA polymerase L

and the ribonucleocapsid template within different halves of the NS phosphoprotein of

vesicular stomatitis virus. Proc. Natl. Acad. Sci. U. S. A. 84, 5655-5659.

27. Finke S, Conzelmann KK, 2003. Dissociation of rabies virus matrix protein functions in

regulation of viral RNA synthesis and virus assembly. J. Virol. 77, 12074-12082.

28. Finke S, Cox JH, Conzelmann KK, 2000. Differential transcription attenuation of rabies virus

genes by intergenic regions: generation of recombinant viruses overexpressing the polymerase

gene. J. Virol. 74, 7261-7269.

29. Schoehn, G, Weissenhorn W, Ruigrok RW, 2006. Crystal structure of the rabies virus

nucleoprotein–RNA complex. Science 313, 360–363.

7

30. Hogenhout SA, Redinbaugh MG, Ammar el D, 2003. Plant and animal rhabdovirus host

range: a bug's view. Trends Microbiol. 11, 264-271.

31. Huang YW, Geng YF, Ying XB, Chen XY, Fang RX, 2005. Identification of a movement

protein of rice yellow stunt rhabdovirus. J. Virol. 79, 2108-2114.

32. Huber LA, Pimplikar S, Parton RG, Virta H, Zerial M, Simons K, 1993. Rab8, a small

GTPase involved in vesicular traffic between the TGN and the basolateral plasma membrane.

J. Cell Biol. 123, 35-45.

33. Hwang LN, Englund N, Das T, Banerjee AK, Pattnaik AK, 1999. Optimal replication activity

of vesicular stomatitis virus RNA polymerase requires phosphorylation of a residue(s) at

carboxy-terminal domain II of its accessory subunit, phosphoprotein P. J. Virol. 73, 5613-

5620.

34. Irie T, Licata JM, Jayakar HR, Whitt MA, Bell P, Harty RN, 2004. Functional analysis of

late-budding domain activity associated with the PSAP motif within the vesicular stomatitis

virus M protein. J. Virol. 78, 7823-7827.

35. IseniF, Barge A, Baudin F, Blondel D, Ruigrok RW, 1998. Characterization of rabies virus

nucleocapsids and recombinant nucleocapsid-like structures. J. Gen. Virol. 79, 2909-2919.

36. Ito N, Moseley GW, Blondel D, Shimizu K, Rowe CL, Ito Y, Masatani T, Nakagawa K, Jans,

DA, Sugiyama M, 2010. Role of interferon antagonist activity of rabies virus phosphoprotein

in viral pathogenicity. J. Virol. 84, 6699-6710.

37. Ito N, Takayama M, Yamada K, Sugiyama M, Minamoto N, 2001. Rescue of rabies virus

from cloned cDNA and identification of the pathogenicity-related gene: glycoprotein gene is

associated with virulence for adult mice. J. Virol. 75, 9121-9128.

38. Ito Y, Nishizono A, Mannen K, Hiramatsu K, Mifune, K, 1996: Rabies virus M protein

expressed in Escherichia coli and its regulatory role in virion-associated transcriptase activity.

Arch. Virol. 141, 671-683.

39. Ivanov I, Yabukarski F, Ruigrok RW, Jamin M, 2011. Structural insights into the rhabdovirus

transcription/replication complex. Virus Res. 162, 126-137.

40. Jackson AO, Dietzgen RG, Goodin MM, Bragg JN, Deng M, 2005. Biology of plant

rhabdoviruses. Ann. Rev. Phytopathol. 43, 623-660.

41. Jacob Y, Badrane H, Ceccaldi PE, Tordo N, 2000. Cytoplasmic dynein LC8 interacts with

lyssavirus phosphoprotein. J. Virol. 74, 10217-10222.

42. Jacob Y, Real E, Tordo N. 2001. Functional interaction map of lyssavirus phosphoprotein:

identification of the minimal transcription domains. J. Virol. 75, 9613-9622.

43. Jayakar HR, Murti KG, Whitt MA, 2003. Mutations in the PPPY motif of vesicular stomatitis

virus matrix protein reduce virus budding by inhibiting a Rabies virus matrix protein regulates

the balance of virus transcription and replication. J. Gen. Virol. 84, 1613-1621.

44. Flamand A, Delagneau JF, 1978. Transcriptional mapping of rabies virus in vivo. J. Virol. 28,

518-523.

45. Flamand A, Raux H, Gaudin Y, Ruigrok RW, 1993. Mechanisms of rabies virus

neutralization. Virology 194, 302-313.

46. Freuling CM, Beer M, Conraths FJ, Finke S, Hoffmann B, Keller B, Kliemt J, Mettenleiter

TC., Muhlbach E, Teifke JP, Wohlsein P, Muller T, 2011. Novel lyssavirus in Natterer's bat,

Germany. Emerg. Infect. Dis. 17, 1519-1522.

47. Gaddy DF, Lyles DS, 2005. Vesicular stomatitis viruses expressing wild-type or mutant M

proteins activate apoptosis through distinct pathways. J. Virol. 79, 4170-4179.

48. Gao Y, Lenard,J, 1995. Multimerization and transcriptional activation of the phosphoprotein

(P) of vesicular stomatitis virus by casein kinase-II. EMBO J. 14, 1240-1247.

49. Gaudin Y, 1997. Folding of rabies virus glycoprotein: epitope acquisition and interaction with

endoplasmic reticulum chaperones. J. Virol. 71, 3742-3750.

50. Gaudin Y, Ruigrok RW, Knossow M, Flamand A, 1993. Low-pH conformational changes of

rabies virus glycoprotein and their role in membrane fusion. J. Virol. 67, 1365-1372.

8

51. Ge P, Tsao J, Schein S, Green TJ, Luo M, Zhou ZH, 2010. Cryo-EM model of the bullet-

shaped vesicular stomatitis virus. Science 327, 689-693.

52. Gholami A, Kassis R, Real E, Delmas O, Guadagnino S, Larrous F, Obach D, Prevost MC,

Jacob Y, Bourhy H, 2008. Mitochondrial dysfunction in lyssavirus-induced apoptosis. J.

Virol. 82, 4774-4784.

53. Lefkowitz EJ (eds), 2006. Virus Taxonomy - 9th Report of the ICTV. Elsevier Academic

Press.

54. Gould AR, Hyatt AD, Lunt R, Kattenbelt JA, Hengstberger S, Blacksell SD, 1998.

Characterisation of a novel lyssavirus isolated from Pteropid bats in Australia. Virus Res. 54,

165-187.

55. Green TJ, Macpherson S, Qiu S, Lebowitz J, Wertz GW, Luo M, 2000. Study of the assembly

of vesicular stomatitis virus N protein: role of the P protein. J. Virol. 74, 9515-9524.

56. Green TJ, Zhang X, Wertz GW, Luo M, 2006. Structure of the vesicular stomatitis virus

nucleoprotein-RNA complex. Science 313, 357-360.

57. Hanlon CA, Kuzmin IV, Blanton JD, Weldon WC, Manangan JS, Rupprecht CE, 2005.

Efficacy of rabies biologics against new lyssaviruses from Eurasia. Virus Res. 111, 44-54.

58. Harty RN, Paragas J, Sudol M, Palese P, 1999. A proline-rich motif within the matrix protein

of vesicular stomatitis virus and rabies virus interacts with WW domains of cellular proteins:

implications for viral budding. J. Virol. 73, 2921-2929.

59. Henry L, Sheff DR, 2008. Rab8 regulates basolateral secretory, but not recycling, traffic at the

recycling endosome. Mol. Biol. Cell 19, 2059-2068.

60. Johal J, Gresty K, Kongsuwan K, Walker PJ, 2008. Antigenic characterization of bovine

ephemeral fever rhabdovirus G and GNS glycoproteins expressed from recombinant

baculoviruses. Arch. Virol. 153, 1657-1665.

61. Johannsdottir HK, Mancini R, Kartenbeck J, Amato L, Helenius A, 2009. Host cell factors

and functions involved in vesicular stomatitis virus entry. J. Virol. 83, 440-453.

62. Kassis R, Larrous F, Estaquier J, Bourhy H, 2004. Lyssavirus matrix protein induces

apoptosis by a TRAIL-dependent mechanism involving caspase-8 activation. J. Virol. 78,

6543-6555.

63. Keene JD, Thornton BJ, Emerson SU, 1881. Sequence-specific contacts between the RNA

polymerase of vesicular stomatitis virus and the leader RNA gene. Proc. Natl. Acad. Sci. U. S.

A. 78, 6191-6195.

64. Kemp GE, Lee VH, Moore DL, Shope RE, Causey OR, Murphy FA. 1973.: Kotonkan, a new

rhabdovirus related to Mokola virus of the rabies serogroup. Am. J. Epidemiol. 98, 43-49.

65. Kim DH, Oh HK, Eou JI, Seo HJ, Kim SK, Oh MJ, Nam SW, Choi TJ, 2005. Complete

nucleotide sequence of the hirame rhabdovirus, a pathogen of marine fish. Virus Res. 107, 1-

9.

66. Knobel DL, Cleaveland S, Coleman PG, Fevre EM, Meltzer MI, Miranda ME, Shaw A,

Zinsstag J, Meslin F.X, 2005. Re-evaluating the burden of rabies in Africa and Asia. Bull.

World Health Organ. 83, 360-368.

67. Kondo H, Maeda T, Shirako Y, Tamada T, 2006. Orchid fleck virus is a rhabdovirus with an

unusual bipartite genome. J. Gen. Virol. 87, 2413-2421.

68. Kopecky SA, Lyles DS, 2003. The cell-rounding activity of the vesicular stomatitis virus

matrix protein is due to the induction of cell death. J. Virol. 77, 5524-5528.

69. Kopecky SA, Lyles DS, 2003. Contrasting effects of matrix protein on apoptosis in HeLa and

BHK cells infected with vesicular stomatitis virus are due to inhibition of host gene

expression. J. Virol. 77, 4658-4669.

70. Kopecky SA, Willingham MC, Lyles DS, 2001. Matrix protein and another viral component

contribute to induction of apoptosis in cells infected with vesicular stomatitis virus. J. Virol.

75, 12169-12181.

9

71. Kouznetzoff A, Buckle M, Tordo N, 1998. Identification of a region of the rabies virus N

protein involved in direct binding to the viral RNA. J. Gen. Virol. 79, 1005-1013.

72. Kretzschmar E, Peluso R, Schnell MJ, Whitt MA, Rose JK, 1996. Normal replication of

vesicular stomatitis virus without C proteins. Virology 216, 309-316.

73. Kurilla MG, Cabradilla CD, Holloway BP, Keene JD, 1984. Nucleotide sequence and host La

protein interactions of rabies virus leader RNA. J. Virol. 50, 773-778.

74. Kuzmin IV, Hughes GJ, Botvinkin AD, Orciari LA, Rupprecht CE, 2005. Phylogenetic

relationships of Irkut and West Caucasian bat viruses within the Lyssavirus genus and

suggested quantitative criteria based on the N gene sequence for lyssavirus genotype

definition. Virus Res. 111, 28-43.

75. Kuzmin IV, Mayer AE, Niezgoda M, Markotter W, Agwanda B, Breiman RF, Rupprecht CE,

2010. Shimoni bat virus, a new representative of the Lyssavirus genus. Virus Res. 149, 197-

210.

76. Kuzmin IV, Novella IS, Dietzgen RG, Padhi A, Rupprecht CE, 2009. The rhabdoviruses:

biodiversity, phylogenetics, and evolution. Infect. Genet. Evol. 9, 541-553.

77. Kuzmin IV, Orciari LA, Arai YT, Smith JS, Hanlon CA, Kameoka Y, Rupprecht CE, 2003.

Bat lyssaviruses (Aravan and Khujand) from Central Asia: phylogenetic relationships

according to N, P and G gene sequences. Virus Res. 97, 65-79.

78. LafayF, Coulon P, Astic L, Saucier D, Riche D, Holley A, Flamand A, 1991. Spread of the

CVS strain of rabies virus and of the avirulent mutant AvO1 along the olfactory pathways of

the mouse after intranasal inoculation. Virology 183, 320-330.

79. Lahaye X, Vidy A, Fouquet B, Blondel D, 2012. Hsp70 protein positively regulates rabies

virus infection. J. Virol. 86, 4743-4751.

80. Lahaye X, Vidy A, Pomier C, Obiang L, Harper F, Gaudin Y, Blondel D, 2009. Functional

characterization of Negri bodies (NBs) in rabies virus-infected cells: Evidence that NBs are

sites of viral transcription and replication. J. Virol. 83, 7948-7958.

81. Langevin C, Jaaro H, Bressanelli S, Fainzilber M, Tuffereau C, 2002. Rabies virus

glycoprotein (RVG) is a trimeric ligand for the N-terminal cysteine-rich domain of the

mammalian p75 neurotrophin receptor. J. Biol. Chem. 277, 37655-37662.

82. Lawson ND, Stillman EA, Whitt MA, Rose JK, 1995. Recombinant vesicular stomatitis

viruses from DNA. Proc. Natl. Acad. Sci. U. S. A. 92, 4477-4481.

83. Le Mercier P, Jacob Y, Tordo N, 1997. The complete Mokola virus genome sequence:

structure of the RNAdependent RNA polymerase. J. Gen. Virol. 78, 1571-1576.

84. Lefrancois L, Lyles DS, 1983. Antigenic determinants of vesicular stomatitis virus: analysis

with antigenic variants. J. Immunol. 130, 394-398.

85. Lentz TL, Burrage TG, Smith AL, Crick J, Tignor GH, 1982. Is the acetylcholine receptor a

rabiesvirus receptor? Science 215, 182-184.

86. Lentz TL, Wilson PT, Hawrot E, Speicher DW, 1984. Amino acid sequence similarity

between rabies virus glycoprotein and snake venom curaremimetic neurotoxins. Science 226,

847-848.

87. Leppert M, Rittenhouse L, Perrault J, Summers DF, Kolakofsky D, 2012. Plus and minus

strand leader pp.183-196.

88. Menager P, Roux P, Megret F, Bourgeois JP, Le Sourd AM, Danckaert A, Lafage M, Prehaud

C, Lafon M. 2009. Toll-like receptor 3 (TLR3) plays a major role in the formation of rabies

virus Negri Bodies. PLoS Pathog 5, e1000315.

89. Mita T, Shimizu K, Ito N, Yamada K, Ito Y, Sugiyama M, Minamoto N, 2008. Amino acid at

position 95 of the matrix protein is a cytopathic determinant of rabies virus. Virus Res. 137,

33-39.

90. Moseley GW, Lahaye X, Roth DM, Oksayan S, Filmer RP, Rowe CL, Blondel D, Jans, DA,

2009. Dual modes of rabies P-protein association with microtubules: a novel strategy to

suppress the antiviral response. J. Cell Sci. 122, 3652-3662.

10

91. Nakahara K, Ohnuma H, Sugita S, Yasuoka K, Nakahara T, Tochikura TS, Kawai A, 1999.

Intracellular behavior of rabies virus matrix protein (M) is determined by the viral

glycoprotein (G). Microbiol. Immunol. 43, 259-270.

92. Nel LH, Rupprecht CE, 2007. Emergence of lyssaviruses in the Old World: the case of Africa.

Curr. Top. Microbiol. Immunol. 315, 161-193.

93. Newcomb WW, Brown JC, 1981. Role of the vesicular stomatitis virus matrix protein in

maintaining the viral nucleocapsid in the condensed form found in native virions. J. Virol. 39,

295-299.

94. Obiang L, Raux H, Ouldali M, Blondel D, Gaudin Y, 2012. Phenotypes of vesicular stomatitis

virus mutants with mutations in the PSAP motif of the matrix protein. J. Gen. Virol. 93, 857-

865.

95. Ogino T, Banerjee AK, 2008. Formation of guanosine(5')tetraphospho(5')adenosine cap

structure by an unconventional mRNA capping enzyme of vesicular stomatitis virus. J. Virol.

82, 7729-7734.

96. Ogino T, Banerjee AK, 2007. Unconventional mechanism of mRNA capping by the RNA-

dependent RNA polymerase of vesicular stomatitis virus. Mol. Cell 25, 85-97.

97. Pasdeloup D, Poisson N, Raux H, Gaudin Y, Ruigrok RW, Blondel D, 2005.

Nucleocytoplasmic shuttling of the rabies virus P protein requires a nuclear localization signal

and a CRM1-dependent nuclear export signal. Virology 334, 284-293.

98. Pattnaik AK, Hwang L, Li T, Englund N, Mathur M, Das T, Banerjee AK, 1997.

Phosphorylation within the amino-terminal acidic domain I of the phosphoprotein of vesicular

stomatitis virus is required for transcription but not for replication. J. Virol. 71, 8167-8175.

99. Peluso RW, Richardson JC, Talon J, Lock M, 1996. Identification of a set of proteins (C' and

C) encoded by the bicistronic P gene of the Indiana serotype of vesicular stomatitis virus and

analysis of their effect on transcription by the viral RNA polymerase. Virology 218, 335-342.

100. Poch O, Blumberg BM, Bougueleret L, Tordo N, 1979.: Sequence comparison of five

polymerases (L proteins) of unsegmented negative-strand RNA viruses: theoretical

assignment of functional domains. J. Gen. RNAs in negative strand virus-infected cells. Cell

18, 735-747.

101. Letchworth GJ, Rodriguez LL, Del Cbarrera J, 2000. Vesicular stomatitis. Vet. J. 157, 239-

260.

102. Lewis P, Fu Y, Lentz TL, 2000. Rabies virus entry at the neuromuscular junction in nerve-

muscle cocultures. Muscle Nerve 23, 720-730.

103. Lichty BD, Power AT, Stojdl DF, Bell JC, 2004. Vesicular stomatitis virus: re-inventing the

bullet. Trends Mol. Med. 10, 210-216.

104. Longdon B, Obbard DJ, Jiggins FM, 2010. Sigma viruses from three species of Drosophila

form a major new clade in the rhabdovirus phylogeny. Proc. Biol. Sci. 277, 35-44.

105. Lyles DS, McKenzie MO, Kaptur PE, Grant KW, Jerome WG, 1996. Complementation of M

gene mutants of vesicular stomatitis virus by plasmid derived M protein converts spherical

extracellular particles into native bullet shapes. Virology 217, 76-87.

106. Marston DA, Ellis RJ, Horton DL, Kuzmin IV,Wise EL, McElhinney LM, Banyard AC,

Ngeleja

107. C, Keyyu J, Cleaveland S, Lembo T, Rupprecht CE., Fooks AR, 2012. Complete genome

sequence of ikoma lyssavirus. J. Virol. 86, 10242-10243.

108. Marston DA, Horton DL, Ngeleja C, Hampson K, McElhinney LM, Banyard AC, Haydon D,

Cleaveland S, Rupprecht CE, Bigambo M, Fooks AR, Lembo T, 2012. Ikoma lyssavirus

(IKOV), highly divergent novel lyssavirus in an African civet. Emerg. Infect. Dis. 18, 664-

667.

109. Masatani T, Ito N, Shimizu K, Ito Y, Nakagawa K, Sawaki Y, Koyama H, Sugiyama M,

2010. Rabies virus nucleoprotein functions to evade activation of the RIG-I-mediated antiviral

response. J. Virol. 84, 4002-4012.

11

110. Mavrakis M, McCarthy AA, Roche S, Blondel D, Ruigrok RW, 2004. Structure and function

of the C-terminal domain of the polymerase cofactor of rabies virus. J. Mol. Biol. 343, 819-

831.

111. McWilliam SM, Kongsuwan K, Cowley JA, Byrne KA, Walker PJ, 1997. Genome

organization and transcription strategy in the complex GNS-L intergenic region of bovine

ephemeral fever rhabdovirus. J. Gen. Virol. 78, 1309-1317.

97. Mebatsion T, Konig M, Conzelmann KK,1996. Budding of rabies virus particles in the

absence of the spike glycoprotein. Cell 84, 941-951.

98. Mebatsion T, Weiland F, Conzelmann KK, 1999. Matrix protein of rabies virus is responsible

for the assembly and budding of bullet-shaped particles and interacts with the transmembrane

spike glycoprotein G. J. Virol. 73, 242-250.

99. Melcher U, 2000. The '30K' superfamily of viral movement proteins. J. Gen. Virol. 81, 257-

266, 2000.

100. Mellon MG, Emerson SU, 1978. Rebinding of transcriptase components (L and NS proteins)

to the nucleocapsid template of vesicular stomatitis virus. J. Virol. 27, 560-567.

101. Rao BL, Basu A, Wairagkar NS, Gore MM, Arankalle VA, Thakare JP, Jadi RS, Rao KA,

Mishra AC, 2003 A large outbreak of acute encephalitis with high fatality rate in children in

Andhra Pradesh, India, in 2003, associated with Chandipura virus. Lancet 364, 869-874.

102. Raux H, Flamand A, Blondel D, 2000. Interaction of the rabies virus P protein with the LC8

dynein light chain. J. Virol. 74, 10212-10216.

103. Reagan KJ, Wunner WH, 1985. Rabies virus interaction with various cell lines is independent

of the acetylcholine receptor. Arch. Virol. 84, 277-282.

104. Rose NF, Publicover J, Chattopadhyay A, Rose JK, 2008. Hybrid alphavirus-rhabdovirus

propagating replicon particles are versatile and potent vaccine vectors. Proc. Natl. Acad. Sci.

U. S. A. 105, 5839-5843.

105. Rupprecht CE, Hanlon CA, Hemachudha T, 2002. Rabies re-examined. Lancet Infect. Dis. 2,

327-343.

106. Sacramento D, Bourhy H, Tordo N, 1991. PCR technique as an alternative method for

diagnosis and molecular epidemiology of rabies virus. Mol. Cell. Probes 5, 229-240.

107. Sato G, Kobayashi Y, Motizuki N, Hirano S, Itou T, Cunha EM, Ito FH, Sakai T, 2009. A

unique substitution at position 333 on the glycoprotein of rabies virus street strains isolated

from non-hematophagous bats in Brazil. Virus Genes 38, 74-79.

108. Schnell MJ, Mebatsion T, Conzelmann KK, 1994. Infectious rabies viruses from cloned

cDNA. EMBO J. 13, 4195-4203.

109. Schoehn G, Iseni F, Mavrakis M, Blondel D, Ruigrok RW, 2001. Structure of recombinant

rabies virus nucleoprotein-RNA complex and identification of the phosphoprotein binding

site. J. Virol. 75, 490-498.

110. Seif I, Coulon P, Rollin PE, Flamand A, 1985. Rabies virulence: effect on pathogenicity and

sequence characterization of rabies virus mutations affecting antigenic site III of the

lycoprotein. J. Virol. 53, 926-934.

111. Shinozaki K, Ebert O, Suriawinata A, Thung SN, Woo SL, 2005. Prophylactic alpha interferon

treatment increases the therapeutic index of oncolytic vesicular stomatitis virus virotherapy for

advanced hepatocellular carcinoma in immune-competent rats. J. Virol. 79, 13705-13713.

112. Sinibaldi L, Goldoni P, Seganti L, Superti F, Tsiang H, Orsi N, 1985. Gangliosides in early

interactions between vesicular stomatitis virus and CER cells. Microbiologica 8, 355-365.

113. Sokol F, Clark HF, 1973. Phosphoproteins, structural components of rhabdoviruses. Virology

52, 246-263.

114. Spiropoulou CF, Nichol ST, 1993. A small highly basic protein is encoded in overlapping

frame within the P gene of vesicular stomatitis virus. J. Virol. 67, 3103-3110.

12

115. Springfeld C, Darai G, Cattaneo R, 2005.: Characterization of the Tupaia rhabdovirus genome

reveals a long open reading frame overlapping with P and a novel gene encoding a small

hydrophobic protein. J. Virol. 79, 6781-6790.

116. St George TD, 1988. Bovine ephemeral fever: a review. Trop. Anim. Health Prod. 20, 194-

202.

117. Stojdl DF, Lichty B, Knowles S, Marius R, Atkins H, Sonenberg N, Bell JC, 2000. Exploiting

tumor-specific defects in the interferon pathway with a previously unknown oncolytic virus.

Nat. Med. 6, 821-825.

118. Superti F, Seganti L, Ruggeri FM, Tinari A, Donelli G, Orsi N, 1987. Entry pathway of

vesicular stomatitis virus into different host cells. J. Gen. Virol.68, 387-399.

119. Takacs AM, Das T, Banerjee AK, 1993. Mapping of interacting domains between the

nucleocapsid protein and the phosphoprotein of vesicular stomatitis virus by using a two-

hybrid system. Proc. Natl. Acad. Sci. U. S. A. 90, 10375-10379.

120. Takayama-Ito M, Inoue K, Shoji Y, Inoue S, Iijima T, Sakai T, Kurane I, Morimoto K, 2006.

A highly attenuated rabies virus HEP-Flury strain reverts to virulent by single amino acid

substitution to arginine at position 333 in glycoprotein. Virus Res. 119, 208-215.

121. Takayama-Ito M, Ito N, Yamada K, Sugiyama M, Minamoto N, 2006. Multiple amino acids

in the glycoprotein of rabies virus are responsible for pathogenicity in adult mice. Virus Res.

115, 169-175.

122. Tan GS, Preuss MA, Williams JC, Schnell MJ, 2007. The dynein light chain 8 binding motif

of rabies virus phosphoprotein promotes efficient viral transcription. Proc. Natl. Acad. Sci. U.

S. A. 104, 7229-7234.

123. Thomas D, Newcomb WW, Brown JC, Wall JS, Hainfeld JF, Trus BL, Steven AC, 1985.

Mass and molecular composition of vesicular stomatitis virus: a scanning transmission

electron microscopy analysis. J. Virol. 54, 598-607.

124. Thomas DC, Roth MG, 1994. The basolateral targeting signal in the cytoplasmic domain of

glycoprotein G from vesicular stomatitis virus resembles a variety of intracellular targeting

motifs related by primary sequence but having diverse targeting activities. J. Biol. Chem. 269,

15732-15739.

125. Thoulouze MI, Lafage M, Schachner M, Hartmann U, Cremer H, Lafon M, 1998. The neural

cell adhesion molecule is a receptor for rabies virus. J.Virol. 72, 7181-7190.

126. Tordo N, Benmansour A, Calisher C, Dietzgen RG, Fang R-X, Jackson AO, Kurath G,

NadinDavis S, Tesh RB, Walker P J, 2005. Family Rhabdoviridae, Fauquet CM, Mayo MA,

Maniloff J, Desselberger U, Ball LA (eds), Virus Taxonomy - 8th Report of the ICTV.

Elsevier Academic Press, London.

127. Walker PJ, Winton JR, 2010. Emerging viral diseases of fish and shrimp. Vet. Res. 41, 51.

128. Wang Y, Walker PJ, 1993. Adelaide river rhabdovirus expresses consecutive glycoprotein

genes as polycistronic mRNAs: new evidence of gene duplication as an evolutionary process.

Virology 195, 719-731.

129. Warrell MJ, Warrell DA, 2004: Rabies and other lyssavirus diseases. Lancet 363, 959-969.

130. Whelan SP, Ball LA, Barr JN, Wertz GT, 1995. Efficient recovery of infectious vesicular

stomatitis virus entirely from cDNA clones. Proc. Natl. Acad. Sci. U. S. A. 92, 8388-8392.

131. Whelan SP, Wertz GW, 1999. The 5' terminal trailer region of vesicular stomatitis virus

contains a position-dependent cis-acting signal for assembly of RNA into infectious particles.

J. Virol. 73, 307-315.

132. Whelan SP, Wertz GW, 1999. Regulation of RNA synthesis by the genomic termini of

vesicular stomatitis virus: identification of distinct sequences essential for transcription but

not replication. J. Virol. 73, 297-306.

133. Whitt MA, Buonocore L, Prehaud C, Rose JK, 1991. Membrane fusion activity, oligomerization,

and assembly of the rabies virus glycoprotein. Virology 185, 681-688.

134. Wu X, Gong X, Foley HD, Schnell MJ, Fu ZF, 2002. Both viral transcription and replication are

13

reduced when the rabies virus nucleoprotein is not phosphorylated. J. Virol. 76, 4153-4161.

135. Wunner WH, 2007. Rabies Virus, Jackson AC e Wunner WH (eds), Rabies, 2nd

ed. Elsevier

Academic Press.

136. Yamada K, Park CH, Noguchi K, Kojima D, Kubo T, Komiya N, Matsumoto T, Mitui MT,

Ahmed K, Morimoto K, Inoue S, Nishizono A, 2012. Serial passage of a street rabies virus in

mouse neuroblastoma cells resulted in attenuation: potential role of the additional N-

glycosylation of a viral glycoprotein in the reduced pathogenicity of street rabies virus. Virus

Res. 165, 34-45.

137. Yang J, Koprowski H, Dietzschold B, Fu ZF, 1999. Phosphorylation of rabies virus nucleoprotein

regulates viral RNA transcription and replication by modulating leader RNA encapsidation. J.

Virol. 73, 1661-1664.

138. Tordo N, Poch O, Ermine A, Keith G, Rougeon F, 1986: Walking along the rabies genome: Is

the large G-L intergenic region a remnant gene? Proc. Natl. Acad. Sci. U. S. A. 83, 3914-3918.

139. Toriumi H, Kawai A, 2004. Association of rabies virus nominal phosphoprotein (P) with viral

nucleocapsid (NC) is enhanced by phosphorylation of the viral nucleoprotein (N). Microbiol.

Immunol. 48, 399-409.

140. Tsiang H, Derer M, Taxi J, 1983.: An in vivo and in vitro study of rabies virus infection of the rat

superior cervical ganglia. Arch. Virol. 76, 231-243.

141. Tuffereau C, Benejean J, Blondel D, Kieffer B, Flamand A, 1998.: Low-affinity nerve-growth

factor receptor (P75NTR) can serve as a receptor for rabies virus. EMBO J. 17, 7250-7259.

142. Tuffereau C, Leblois H, Benejean J, Coulon P, Lafay F, Flamand A, 1989. Arginine or lysine in

position 333 of ERA and CVS glycoprotein is necessary for rabies virulence in adult mice.

Virology 172, 206-212.

143. Vandepol SB, Lefrancois L, Holland JJ, 1986. Sequences of the major antibody binding epitopes of

the Indiana serotype of vesicular stomatitis virus. Virology 148, 312-325.

144. Vidy A, Chelbi-Alix M, Blondel D, 2005. Rabies virus P protein interacts with STAT1 and

inhibits interferon signal transduction pathways. J. Virol. 79, 14411-14420.

145. Vidy A, El Bougrini J, Chelbi-Alix MK, Blondel D, 2007. The nucleocytoplasmic rabies virus

P protein counteracts interferon signaling by inhibiting both nuclear accumulation and DNA

binding of STAT1. J. Virol. 81, 4255-4263.

146. von Kobbe C, van Deursen JM, Rodrigues JP, Sitterlin D, Bachi A, Wu X, Wilm M, Carmo-

Fonseca M, Izaurralde E, 2000. Vesicular stomatitis virus matrix protein inhibits host cell gene

expression by targeting the nucleoporin Nup98. Mol. Cell 6, 1243-1252.

147. Walker PJ, Byrne KA, Cybinski DH, Doolan DL, Wang YH, 1991. Proteins of bovine

ephemeral fever virus. J. Gen. Virol. 72, 67-74.

151�Walker PJ, Byrne KA, Riding GA, Cowley JA,Wang Y, McWilliam S, 1992. The genome of

bovine ephemeral fever rhabdovirus contains two related glycoprotein genes. Virology 191,

49-61.