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Descriptions and Lists from the VIDE Database

Cucumber mosaic cucumovirus

Index

• Nomenclature
• Host range, transmission and symptoms
• Physical and biochemical properties
• Taxonomy and relationships
• Comments and References

Data collated by R.I.B. Francki, 1980; N. Habili, 1987.

Nomenclature

Synonyms

banana infectious chlorosis virus, coleus mosaic virus (Creager, 1945; Holcomb and Valverde, 1991), cowpea banding mosaic virus, cowpea ringspot virus, cucumber virus 1, lily ringspot virus (Brierley and Travis, 1958), pea top necrosis virus, peanut yellow mosaic virus, southern celery mosaic virus (Doolittle, 1916; Price, 1935; Wellman, 1934), soybean stunt virus (Hanada and Tochihara, 1982), spinach blight virus, tomato fern leaf virus, pea western ringspot virus.

Acronym

CMV

Strains

numerous, the better known include: A-CMV, E-CMV, L-CMV, N-CMV, P-CMV, Z-CMV and WAI/WAII; there seem to be two antigenic groups, ToRS and DTL (Devergne, 1975).

ICTV decimal code

10.0.4.0.001

Host range and symptoms

Natural host range and symptoms

Symptoms persist.
• Very many hosts (Kaper and Waterworth, 1981) including:
• Cucumis sativus and many other cucurbits - mosaics and stunting, reduced fruit yield.
• Lycopersicon esculentum - mosaic, reduction of leaf laminae ("fernleaf") and stunting.
• Spinacia oleracea - severe chlorosis and stunting.

Transmission

Transmitted by a vector; an insect; more than 60 spp. including Acyrthosiphon pisum, Aphis craccivora and Myzus persicae; Aphididae. Transmitted in a non-persistent manner. Virus transmitted by mechanical inoculation; transmitted by seed (in 19 species, but in variable extents).

Geographical distribution

Probably distributed worldwide.

Experimental host range

Many (>9) families susceptible.

Diagnostically susceptible host species and symptoms

• Chenopodium amaranticolor, C. quinoa - chlorotic local lesions.
• Cucumis sativus - systemic mosaic.
• Vigna unguiculata - necrotic local lesions.
• Lycopersicon esculentum, Nicotiana × edwardsonii, N. glutinosa, N. tabacum - symptoms depend on virus strain.

Maintenance and propagation hosts

Cucumis sativus, Nicotiana clevelandii, N. glutinosa, N. tabacum.

Assay hosts (Local lesions or Whole plants)

Chenopodium amaranticolor (L), C. quinoa (L), Vigna unguiculata (L).

Susceptible host species

• Abelmoschus esculentus
• Amaranthus caudatus
• Amaranthus retroflexus
• Antirrhinum majus
• Apium graveolens
• Atriplex hortensis
• Beta vulgaris
• Brassica campestris ssp. napus
• Brassica campestris ssp. pekinensis
• Brassica campestris ssp. rapa
• Brassica juncea
• Calendula officinalis
• Capsella bursa-pastoris
• Capsicum annuum
• Capsicum frutescens
• Catharanthus roseus
• Cheiranthus cheiri
• Chenopodium album
• Chenopodium amaranticolor
• Chenopodium foetidum
• Chenopodium hybridum
• Chenopodium murale
• Chenopodium quinoa
• Chrysanthemum morifolium
• Cicer arietinum
• Cichorium endiva
• Citrullus lanatus
• Coriandrum sativum
• Crotalaria spectabilis
• Cucumis melo
• Cucumis sativus
• Cucurbita maxima
• Cucurbita moschata
• Cucurbita pepo
• Datura ferox
• Datura metel
• Datura stramonium
• Datura tatula
• Daucus carota
• Emilia sagittata
• Fagopyrum esculentum
• Glycine max
• Gomphrena globosa
• Gossypium hirsutum
• Helianthus annuus
• Hyoscyamus niger
• Ipomoea nil
• Lens culinaris
• Lotus corniculatus
• Lupinus albus
• Lupinus angustifolius
• Lycopersicon esculentum
• Lycopersicon pimpinellifolium
• Matthiola incana
• Medicago sativa
• Melilotus albus
• Momordica balsamina
• Nicotiana bigelovii
• Nicotiana clevelandii
• Nicotiana debneyi
• Nicotiana glutinosa
• Nicotiana megalosiphon
• Nicotiana rustica
• Nicotiana sylvestris
• Nicotiana tabacum
• Nicotiana × edwardsonii
• Phaseolus lunatus
• Phaseolus vulgaris
• Physalis floridana
• Physalis peruviana
• Phytolacca americana
• Pisum sativum
• Raphanus sativus
• Rumex acetosa
• Senecio vulgaris
• Solanum demissum
• Solanum melongena
• Solanum nigrum
• Solanum nodiflorum
• Solanum rostratum
• Solanum tuberosum
• Sonchus oleraceus
• Spinacia oleracea
• Tetragonia tetragonioides
• Trifolium hybridum
• Trifolium incarnatum
• Trifolium pratense
• Trifolium repens
• Tropaeolum majus
• Verbesina encelioides
• Vicia faba
• Vicia sativa
• Vigna radiata
• Vigna unguiculata
• Vigna unguiculata ssp. sesquipedalis

Insusceptible host species

• Celosia argentea
• Citrullus vulgaris
• Dahlia pinnata
• Lactuca sativa
• Lathyrus odoratus
• Nicandra physalodes
• Petunia × hybrida
• Spinacia oleracea
• Stellaria media
• Zinnia elegans

Families containing susceptible hosts

• Amaranthaceae (3/4)
• Apocynaceae (1/1)
• Chenopodiaceae (9/9)
• Compositae (8/11)
• Convolvulaceae (1/1)
• Cruciferae (8/8)
• Cucurbitaceae (7/8)
• Leguminosae-Papilionoideae (21/22)
• Malvaceae (2 /2)
• Phytolaccaceae (1/1)
• Polygonaceae (2/2)
• Scrophulariaceae (1/1)
• Solanaceae (26/28)
• Tetragoniaceae (1/1)
• Tropaeolaceae (1/1)
• Umbelliferae (3/3)

Families containing insusceptible hosts

• Amaranthaceae (1/4)
• Caryophyllaceae (1/1)
• Chenopodiaceae (1/9)
• Compositae (3/11)
• Cucurbitaceae (1/8)
• Leguminosae-Papilionoideae (1/22)
• Solanaceae (2/28)

Sources of host-range data

Douine et al. (1979).

Physical and biochemical properties

Properties of particles in sap

TIP: 55-70 °C. LIV: 1-10 days. DEP: log₁₀ minus 3-6. Infectivity of sap not changed by treatment with di-ethyl ether. Leaf sap contains few virions, or contains many virions. Electron microscopy: freeze-dry preparations for shadowing, or fix with aldehyde for negative staining with PTA.

Purification method

Lot et al. (1972) and modified by Peden and Symons in 1973 (Clark et al., 1974).

Particle morphology

Virions isometric; not enveloped; 29 nm in diameter; rounded in profile; without a conspicuous capsomere arrangement.

Physical properties

One sedimenting component in purified preparations; sedimentation coefficient 99 S. Density 1.367 g cm^-3 in CsCl (after fixation with formaldehyde). Isoelectric point pH 5.5 (Q strain). A₂₆₀/A₂₈₀ ratio 1.7 (corrected).

Biochemical properties

Virions contain 18 % nucleic acid; 82 % protein; 0 % lipid.

Genome consists of RNA; single-stranded; linear. Total genome size 8.621 kb. Genome of three parts; largest (or only) genome part the largest 3.389 kb; the 2nd largest 3.035 kb; the 3rd largest 2.197 kb. Genomic nucleic acid isolated by Gould and Symons (1977). Base composition 24 % G; 23 % A; 23 % C; 30 % U. 5´ terminus of RNA has a methylated nucleotide cap. Infectivity retained when deproteinised with proteases, or decreased when deproteinised with proteases; retained when deproteinised with phenol or detergent. Poly A region absent. Genome has tRNA-like activity. Genome accepts tyrosine.

NCBI sequence data (Entrez search)

Sequence database accession code(s)

• D00355 Em(40)_vi:CUCMVRN2 Gb(84)_vi:MCVRN2 Cucumber mosaic Virus Fny strain RNA2. 8/90 3,050bp.
• D00356 Em(40)_vi:CUMCVFRN Gb(84)_vi:MCVFRNA1 (Fny strain) RNA1. 3/91 3,357bp.
• D00385 Em(40)_vi:CUMCVO3 Gb(84)_vi:MCVO3, strain O, RNA 3, virus genome. 4/90 2,217bp.
• D00462 Em(40)_vi:MCVRNA3C Gb(84)_vi:MCVRNA3C RNA 3, coat protein gene and flanks. 3/91 1,255bp.
• D00463 Em(40)_vi:MCVRNA3W Gb(84)_vi:MCVRNA3WL RNA 3, coat protein gene and flanks. 3/91 1,267bp.
• D10209 Em(40)_vi:MCVORNA2 Gb(84)_vi:MCVORNA2 (CMV) RNA 2. 10/92 3,049bp.
• D10538 Em(40)_vi:MCVCP3A1 Gb(84)_vi:MCVCP3A1 (Fny-CMV) RNA 3, complete genome. 6/92 2,216bp.
• D10539 Em(40)_vi:MCVCP3A2 Gb(84)_vi:MCVCP3A2 (M-CMV) RNA 3, complete genome. 6/92 2,215bp.
• D10544 Em(40)_vi:MCVCPFC Gb(84)_vi:MCVCPFC (CMV-FC) coat protein, complete cds. 6/92 657bp.
• D10545 Em(40)_vi:MCVCPP6 Gb(84)_vi:MCVCPP6 (CMV-P6) coat protein, complete cds. 6/92 657bp.
• D12499 Em(40)_vi:MCVYRNA3 Gb(84)_vi:MCVYRNA3 RNA3, 3a protein and coat protein genes, complete cds. 9/92 2,217bp.
• D12537 Em(40)_vi:MCVRNA1 Gb(84)_vi:D12537 RNA1, complete sequence. 10/92 3,361bp.
• D12538 Em(40)_vi:MCVRNA2 Gb(84)_vi:D12538 RNA2, complete sequence. 10/92 3,051bp.
• D16403 Em(40)_vi:MCVSVR1 Gb(84)_vi:MCVL1 RNA for viral replicase subunit, complete cds. 11/93 3,359bp
• D16405 Em(40)_vi:MCVSVR2 Gb(84)_vi:MCVL3 RNAs for movement protein and viral coat protein, complete cds. 11/93 2
• D16406 Em(40)_vi:MCVPMPVCP Gb(84)_vi:MCVL2 RNA for viral replicase subunit, complete cds. 11/93 3,047bp
• D28778 Em(40)_vi:MCV1A1 Gb(84)_vi:MCV1A1 RNA for 1a. 3/94 3,358bp.
• D28779 Em(40)_vi:MCV2A2 Gb(84)_vi:MCV2A2 RNA for 2a and 2b. 3/94 3,042bp.
• D28780 Em(40)_vi:MCV3A3 Gb(84)_vi:MCV3A3 RNA for 3a and coat protein. 3/94 2,214bp
• D42079 Em(43)_vi:Mcvr3mpcp Gb(89)_vi:Mcvr3mpcp1 Cucumber mosaic virus gene for movement protein and viral coat protein. 11/94 2,238bp.
• D42080 Em(43)_vi:Mcvr3mpc1 Gb(89)_vi:Mcvr3mpcp2 Cucumber mosaic virus gene for movement protein and viral coat protein. 11/94 2,214bp.
• D42081 Em(43)_vi:Mcvst3acp Gb(89)_vi:Mcvst3acp Cucumber mosaic virus gene for movement protein and coat protein. 11/94 334bp.
• D42082 Em(43)_vi:Mcvsrmpcp Gb(89)_vi:Mcvsrmpcpa Cucumber mosaic virus gene for movement potein and coat protein. 11/94 334bp.
• D43798 Em(43)_vi:Mcv3apa Gb(89)_vi:Mcv3apa Cucumber mosaic virus mRNA for 3a protein. 12/94 840bp.
• D43799 Em(43)_vi:Mcvr1pb Gb(89)_vi:Mcvr1pb Cucumber mosaic virus mRNA for RNA1 protein. 12/94 1,137bp.
• D43800 Em(43)_vi:Mcvcp Gb(89)_vi:Mcvcp Cucumber mosaic virus mRNA for coat protein. 12/94 733bp.
• D49496 Em(43)_vi:Mcv3acp Gb(89)_vi:Mcv3acp Cucumber mosaic virus gene for 3a and cp. 3/95 2,205bp.
• J02059 Em(40)_vi:MCVRNA3 Gb(84)_vi:MCVRNA3 (Q strain) RNA 3, complete segment including RNA 4 cds. 5/92 2,193bp.
• K01781 Gb(84)_vi:MCVRNA1 (Q strain) RNA 1, 3´ end. 1/85 271bp.
• K01782 Gb(84)_vi:MCVRNA2 (Q strain) RNA 2, 3´ end. 1/85 271bp.
• L15336 Em(40)_vi:MCV3APCOA Gb(84)_vi:MCV3APCOAT 3A protein and coat protein mRNA (RNA3) complete cds. 5/93 2,209bp.
• L36251 Em(43)_vi:Mcvrna3ko Gb(89)_vi:Mcvrna3kor Cucumber mosaic virus RNA3 encoding movement protein (MP) and capsid protein (CP) genes.
• L36525 Em(43)_vi:Mcvcpb Gb(89)_vi:Mcvcpb Cucumber mosaic virus coat protein (CP) gene, complete cds. 10/94 894bp.
• L40953 Em(43)_vi:Mcvcopra Gb(89)_vi:Mcvcopra Cucumber mosaic virus coat protein, complete cds. 4/95 757bp.
• M21464 Em(40)_vi:CUMCVRNA Gb(84)_vi:MCVRNA3A RNA 3 containing 3A protein and coat protein, complete cds. 7/89 2,197b
• M22710 Em(40)_vi:CUMCVCPA Gb(84)_vi:MCVCPA coat protein gene, complete cds. 4/90 1,362bp.
• M29129 Em(40)_vi:CUMCVCOP Gb(84)_vi:MCVCOPR coat protein (segment 4) mRNA, partial cds. 2/90 106bp.
• M57602 Em(40)_vi:MCVYCNS Gb(84)_vi:MCVYCNS Cucumber mosaic virus Y strain, 3A protein, coat protein mRNA, complete cds. 9/92 2,217bp.
• M98499 Em(40)_vi:MCVCPAA Gb(84)_vi:MCVCPAA coat protein (CP) RNA, complete cds. 8/92 657bp.
• M98500 Em(40)_vi:MCVCPAB Gb(84)_vi:MCVCPAB coat protein (CP) RNA, complete cds. 8/92 657bp.
• M98501 Em(40)_vi:MCVCPAC Gb(84)_vi:MCVCPAC coat protein (CP) RNA, complete cds. 8/92 657bp.
• S70105 Em(40)_un:S70105 Gb(84)_vi:S70105 cp=coat protein CMV, subgroup II, WL,Genomic, 1360 nt. 2/93 1,360bp.
• S72187 Gb(89)_un:S72187 orf2a, orf2b (cucumber mosaic cucumovirus CMV, K, Genomic RNA, nt). 10/94 3,048bp.
• U10922 Gb(84)_vi:CMU10922 DKRD coat protein gene, complete cds. 6/94 834bp.
• U10923 Gb(84)_vi:CMU10923 SP103 coat protein gene, complete cds. 6/94 841bp.
• U10924 Gb(84)_vi:CMU10924 SP104 coat protein gene, complete cds. 6/94 835bp.
• U20668 Em(43)_vi:Cm20668 Gb(89)_vi:Cmu20668 Cucumber mosaic virus truncated protein 3a and coat protein mRNA, complete cds. 5/95 1,906bp.
• U22821 Em(43)_vi:Cm22821 Gb(89)_vi:Cmu22821 Cucumber mosaic virus Ny coat protein gene, complete cds. 4/95 1,033bp.
• U22822 Em(43)_vi:Cm22822 Gb(89)_vi:Cmu22822 Cucumber mosaic virus Sn coat protein gene, complete cds. 4/95 1,041bp.
• U31219 Em(44)n:Cm31219 Gb(90)n:Cmu31219 Cucumber mosaic virus Hawaii coat protein mRNA, complete cds. 8/95 657bp.
• U31220 Em(44)n:Cm31220 Gb(90)n:Cmu31220 Cucumber mosaic virus Oahu coat protein mRNA, complete cds. 8/95 657bp.
• U32858 Em(44)n:Cm32858 Gb(90)n:Cmu32858 Cucumber mosaic virus, from plantains, coat protein (CP) mRNA, complete cds. 8/95 864bp.
• U32859 Em(44)n:Cm32859 Gb(90)n:Cmu32859 Cucumber mosaic virus, from bananas, coat protein (CP) mRNA, complete cds. 8/95 862bp.
• X00817 Em(40)_vi:CURNA01 Gb(84)_vi:CURNA01 strain Y (CMV-Y) RNA 5 sequence. 5/92 368bp.
• X00985 Em(40)_vi:CVRNA02 Gb(84)_vi:CVRNA02 RNA 2 (Q-CMV RNA 2). 9/93 3,035bp.
• X01333 Em(40)_vi:CURNA1 Gb(84)_vi:CURNA1 (CMV) RNA1 3´ terminal region. 11/85 270bp.
• X01334 Em(40)_vi:CURNA2 Gb(84)_vi:CURNA2 (CMV) RNA2 3´ terminal region. 5/92 270bp
• X01335 Em(40)_vi:CURNA3 Gb(84)_vi:CURNA3 (CMV) RNA3 3´ terminal region. 11/85 267bp.
• X01336 Em(40)_vi:CURNA4 Gb(84)_vi:CURNA4 (CMV) RNA4 3´ terminal region. 11/85 267bp.
• X02733 Em(40)_vi:CURNA1Q Gb(84)_vi:CURNA1Q strain Q (Q-CMV) RNA 1 sequence. 9/93 3,389bp.
• X16386 Em(40)_vi:CUMCVRN4 Gb(84)_vi:CUMCVRN4 (CMV) RNA4 gene for coat protein. 9/93 1,007bp.
• X65017 Em(40)_vi:CMVCOPR Gb(84)_vi:CMVCOPR genomic RNA for coat protein. 6/92 963bp.
• X77855 Em(40)_vi:CMVCOATP Gb(84)_vi:CMVCOATP (As) RNA4 mRNA for coat protein. 2/94 1,043bp.
• X81161 Em(43)_vi:Cmvt8920 Gb(89)_vi:Cmvt8920 Cucumber mosaic Virus T/89/20 genomic RNA. 8/94 450bp.
• X81162 Em(43)_vi:Cmvt8943 Gb(89)_vi:Cmvt8943 Cucumber mosaic Virus T/89/43 genomic RNA. 8/94 450bp.
• X81163 Em(43)_vi:Cmvt9020 Gb(89)_vi:Cmvt9020 Cucumber mosaic Virus T/90/20 genomic RNA. 8/94 450bp.
• X81164 Em(43)_vi:Cmvt9023 Gb(89)_vi:Cmvt9023 Cucumber mosaic Virus T/90/23 genomic RNA. 8/94 450bp.
• X81165 Em(43)_vi:Cmvt9104 Gb(89)_vi:Cmvt9104 Cucumber mosaic Virus T/91/04 genomic RNA. 8/94 450bp.
• X81166 Em(43)_vi:Cmvt9102 Gb(89)_vi:Cmvt9102 Cucumber mosaic Virus T/91/02 genomic RNA. 8/94 450bp.
• X81167 Em(43)_vi:Cmvt9107 Gb(89)_vi:Cmvt9107 Cucumber mosaic Virus T/91/07 genomic RNA. 8/94 450bp.
• X81168 Em(43)_vi:Cmvt9109 Gb(89)_vi:Cmvt9109 Cucumber mosaic Virus T/91/09 genomic RNA. 8/94 450bp.
• X81169 Em(43)_vi:Cmvt9224 Gb(89)_vi:Cmvt9224 Cucumber mosaic Virus T/92/24 genomic RNA. 8/94 450bp.
• X81170 Em(43)_vi:Cmv920 Gb(89)_vi:Cmv920 Cucumber mosaic Virus T/92/09 genomic RNA. 8/94 450bp.
• X89652 Em(44)n:Cmvcpgene Gb(90)_vi:Cmvcpgene Cucumber Mosaic Virus (CMV) RNA for capsid protein gene. 7/95 880bp.
• Z12818 Em(40)_vi:CMV3ACP Gb(84)_vi:CMV3ACP genes for 3a protein and coat protein. 2/94 2,199bp.
• Z21863 Em(40)_vi:CM2BPRO Gb(84)_vi:CM2BPRO RNA 4A. 3/94 682bp.

Features of the genome

Non-genomic nucleic acid found in the virions; is subgenomic mRNA (of coat protein), or satellite RNA (in some isolates). Sub-genomic mRNA found in infected cells; coat protein mRNA, which is also the encapsidated RNA-4 of 1.027 kb.

Features of proteins

Virion protein(s) one; M_r 24247; coat protein. Method of preparation: Habili and Francki (1974). Amino acid sequence: Davies and Symons (1987). Amino acid composition: Mossop (1978).

Virus-coded non-virion proteins identified by genomic sequence analysis (Clark et al. (1974); Kumarasamy and Symons (1979); Gill et al. (1981)); three proteins found. M_r of the largest 36300 (791 residues); RNA-1 translation product from Q-CMV, putative viral RNA replicase component (Rezaian et al., 1985). M_r of 2nd largest 31000 (333 residues; RNA-2 translation product from Q-CMV, putative viral RNA polymerase (Rezaian et al., 1985)). M_r of 3rd 9900 (353 residues); 3a protein, possible movement protein (Davies and Symons, 1987).

Replication

Replication does not depend on a helper virus.

Cytopathology

Virions found in all parts of the host plant; in cytoplasm. Inclusions present in infected cells; are crystals in the cytoplasm (that are often rhomboidal, hexagonal or roughly spherical and may appear as solid hollow structures); they contain virions.

Taxonomy and relationships

Cucumovirus: Bromoviridae (the type species)

Virus(es) with serologically related virions

Peanut stunt and tomato aspermy viruses, but distantly.

Additional comments on relationships

Sequence homologies indicate that the virus is distantly related to brome mosaic virus, and also, in sequences in RNA-1 and RNA-2, to alfalfa mosaic and tobacco mosaic viruses (Rezaian et al., 1985; Davies and Symons, 1987).

Comments and References

General comments

Pea western ringspot virus is probably a mixture of cucumber mosaic and clover yellow vein viruses.

References

• Brierley, P. and Travis, R.V. (1958). Pl. Dis. Reptr 42: 1034.
• Clark, G.L., Peden, K.W.C. and Symons, R.H. (1974). Virology 62: 434.
• Creager, D.B. (1945). Phytopathology 35: 223.
• Devergne, J.-C. (1975). Meded. Facult. Landb. Rijks-Univ. Gent 40: 19.
• Doolittle, S.P. (1916). Phytopathology 6: 145.
• Douine, L., Quiot, J.B., Marchoux, G. and Archange, P. (1979). Annls. Phytopath. 11: 439.
• Finch, J.T., Klug, A. and van Regenmortel, M.H.V. (1967). J. mol. Biol. 24: 303.
• Francki, R.I.B., Mossop, D.W. and Hatta, T. (1979). CMI/AAB Descr. Pl. Viruses No. 213, 6 pp.
• Gibbs, A.J. and Harrison, B.D. (1970). CMI/AAB Descr. Pl. Viruses No. 1, 4 pp.
• Gill, D.S., Kumarasamy, R. and Symons, R.H. (1981). Virology 113: 1.
• Habili, N. and Francki, R.I.B. (1974). Virology 57: 392.
• Hanada, K. and Tochihara, H. (1982). Phytopathology 72: 761.
• Holcomb, G.E. and Valverde, R.A. (1991). Plant Dis. 75: 1183.
• Jagger, I.C. (1916). Phytopathology 6: 148.
• Kaper, J.M. and Waterworth, H.C. (1981). In: Plant Virus Infections: Comparative Diagnosis, pp.257-332; ed. E. Kurstak. Elsevier/North Holland, Amsterdam.
• Komuro, Y. (1973). Viruses in Vegetables. Seibundo Shinkosha, Tokyo.
• Kumarasamy, R. and Symons, R.H. (1979). Virology 96: 622.
• Lot, H., Jonard, G. and Richards, K. (1977). FEBS Lett. 80: 395.
• Mossop, D.W. (1978). Ph.D. Thesis, University of Adelaide, Australia.
• Price, W.C. (1934). Phytopathology 24: 743.
• Price, W.C. (1935). Phytopathology 25: 947.
• Rezaian, M.A., Williams, R.H.V. and Symons, R.H. (1985). Eur. J. Biochem. 150: 331.
• Wellman, F.L. (1934). Phytopathology 24: 695.

Illustrations

Cite this publication as: Brunt, A.A., Crabtree, K., Dallwitz, M.J., Gibbs, A.J., Watson, L. and Zurcher, E.J. (eds.) (1996 onwards). `Plant Viruses Online: Descriptions and Lists from the VIDE Database. Version: 20^th August 1996.' URL http://biology.anu.edu.au/Groups/MES/vide/

Dallwitz (1980) and Dallwitz, Paine and Zurcher (1993) should also be cited.