Tobacco mosaic virus (type strain)
Dept. of Plant Pathology, Cornell University, Ithaca, New York 14853, USA
H. W. Israel
Dept. of Plant Pathology, Cornell University, Ithaca, New York 14853, USA
Disease described by
Iwanowski (1892) and
description relates more to recent isolates of the virus for which chemical, physical
and biological measurements have been made. Exact relationships of these isolates to the
original is not known, but from the following it is presumed that all type
very similar, if not identical. Virus obtained from W. M. Stanley
independently sub-cultured many times in the USA and Germany, and some 20 years later
its coat protein was shown in both countries to have the same amino-acid sequence.
- Common strain, wild type, ordinary TMV, vulgare (Rev. appl. Mycol. 23:
- U1 (Rev. appl. Mycol. 34: 110)
- Marmor tabaci (Rev. appl. Mycol. 28: 514)
An RNA-containing virus with rigid, tubular, helically symmetrical particles c.
18 x 300 nm. Wide host range, world-wide distribution. Easily transmitted by
mechanical inoculation, but not by insects or other common vectors. The most
investigated plant virus; important more for basic research than as an agent of
economically important diseases.
Elicits disease in many plant species
although exact strain not always
described. Causes economically important mosaic diseases in tobacco (Nicotiana
Common in countries where tobacco is grown in the field.
Host Range and SymptomatologyHolmes (1946)
obtained infection in 199 of 310 species from 30 families of
angiosperms, although in 100 of these only the inoculated leaves became infected.
Species differ in their capacity to support virus replication; many, including ferns
support only very small levels
(Cheo & Girard, 1971
- Nicotiana tabacum cvs. Turkish, Turkish Samsun, Samsun (Samsoun), White Burley,
Burley and Xanthi. Vein-clearing appears on young, systemically-invaded leaves 3-4 days
after inoculation, followed by a light green-dark green mosaic
by distortion and blistering. Inoculated leaves exhibit few symptoms other than faint
chlorotic lesions when the nutrient nitrogen supply is limited.
- N. glutinosa, N. tabacum cvs. Samsun NN and Xanthi-nc, Phaseolus vulgaris
cv. Pinto and Chenopodium amaranticolor form necrotic local lesions at
temperatures below c. 28°C; systemic infections appear in the Nicotiana
spp. at higher temperatures.
- N. sylvestris, and N. tabacum cv. Java develop systemic infections;
however, strains or mutants may produce necrotic local lesions without systemic infection.
- N. tabacum cvs. Turkish, Turkish Samsun, Samsun or Xanthi.
- Local lesion assays are most frequently performed with N. tabacum cvs.
Xanthi-nc or Samsun NN, N. glutinosa, Phaseolus vulgaris cv. Pinto or Chenopodium
StrainsHennig & Wittmann (1972)
list many well-documented
the subject of other Descriptions in this series. Tobacco plants infected with type
strain may give rise to variants which can be isolated from
yellow or necrotic spots
in systemically-invaded tissue. A masked strain inducing no symptoms in Samsun
tobacco has been selected by growing type strain-infected plants at 35°C
Many chemically induced mutants have been isolated
(Hennig & Wittmann, 1972
Transmission by Vectors
Considered not normally transmissible by arthropods but is readily spread between
plants by contact and by man during cultural operations. Scattered reports of insects
(Lojek & Orlob, 1969
Harris & Bradley, 1973
relate to their acting
inadvertently as agents of mechanical inoculation. Soil-borne virus particles or fragments
of infected tissue can serve as sources of infection via roots.
Transmission through Seed
Type strain not transmissible by seed or pollen.
Transmission by Dodder
Transmitted, chiefly among tobaccos, by Cuscuta campestris, C. japonica
but the specific virus strains have not always
Good immunogen; high titred antisera can be obtained with either whole virus or
coat protein subunits. Easily detected by interfacial ring precipitation tests,
quantitative precipitin tests, radioimmunoassay, inhibition of infectivity by serum,
etc., and by a variety of gel-diffusion procedures
with intact virus develop only after several days because the antigen is large. Topic
van Regenmortel (1966)
Strains and mutants exhibit differing amounts of serological cross reactivity with
type strain, depending on the nature and position of amino acid replacements in the coat
(von Sengbusch, 1965
van Regenmortel, 1967
Some symptom variants, including
the masked strain, seem serologically identical with the type strain. Relationships to
isolates are being dealt with in a separate
Stability in Sap
Very stable; preparations retain infectivity for decades
(Silber & Burk, 1965
Also very heat-stable; some infectivity is retained after 10 min exposures at over
90°C. Dilutions of 10-6
of expressed tobacco sap can be infectious.
Because of its high titre and stability, TMV can be purified by many procedures such
as ultracentrifugation, or salt, isoelectric or solvent precipitations. The polyethylene
glycol procedure of
Gooding & Hebert (1967)
also yields satisfactory preparations;
modifications include the use of buffered 5% Triton X-100 to eliminate chloroplast
(Nozu & Yamura, 1971
and incubation of virus with chelating agents to
remove coloured materials
(Ginoza et al., 1954
Resuspension of sedimented virus
in dilute ethylenediamine-tetraacetate (EDTA) reduces virus aggregation
(Boedtker & Simmons, 1958
Particles can be sorted according to length using columns of agar or
Yields may reach 10 mg/g fresh weight, but 1-3 are more
Properties of Particles
Sedimentation coefficient (s20, w
) at infinite dilution is c.
(Harrington & Schachman, 1956
M. Wt is c. 39.4 x
Diffusion coefficient (D20,w)
is c. 4.4 x 10-8 cm2/sec
(Schramm & Bergold, 1947).
Isoelectric point is c. pH 3.5
(Fraenkel-Conrat & Narita, 1958).
specific volume is c. 0.73 cm3/gm
mobility at ionic strength (G) 0.075 and pH 6.5-7.9 is
c. -0.83 (µm/sec)/(V/cm)
(Kramer & Wittmann, 1958).
coefficient (A(0.1%,1 cm), at 260 nm, uncorrected for light scattering, ranges
between 2.7 and 3.5
a value of 3.0 is commonly used.
A260/A280 is c. 1.19
Buoyant density in CsCl is c. 1.325
(Siegel & Hudson, 1959).
Straight, rigid tubules; length c.
300 nm, max. radius c.
composed of c.
2130 identical protein subunits closely packed in a helix
2.3 nm, 16 1/3 subunits/turn) around a cylindrical canal of radius
2 nm. One continuous single strand of RNA, of c.
follows the same helix (49 nucleotides/turn or 3/subunit) at a radius of c.
4 nm, and is associated with the protein subunits near their inner surfaces
Particles can be dissociated into constituent nucleic
acid and protein and reconstituted into stable infective viral particles
Particle CompositionNucleic acid.
RNA, single-stranded, M. Wt 2.05 x 106
5% of particle weight. G:A:C:U = 25.3:29.8:18.5:26.3
subsidiary RNA components in virus particle, but pieces of viral RNA occur in tissues
(Siegel et al., 1973
Preparation of infective RNA described by
Mandeles & Bruening (1968)
is 32.5 S in 0.05 M KCl, 2 mM EDTA, 5
µg/ml sodium dodecyl sulphate.
Protein: Can be isolated according to
About 95% of
particle; subunits, of M. Wt c. 1.75 x 104, each consist of a chain
of 158 amino acids with the following sequence
(Wittmann-Liebold & Wittmann, 1967):
Note: In the Japanese common strain (OM) the threonine at position 153 is replaced
by asparagine and the isoleucine at position 129 is replaced by valine
(Nozu et al., 1970).
Particles have no known enzymatic activity, but RNase is a troublesome contaminant
(Whitfeld & Williams, 1963).
Other components: Traces of metal
(Loring et al., 1962)
and a polyamine
(Johnson & Markham, 1962),
the nature of which is challenged
(Beer & Kosuge, 1970),
are reported as possible constituents of the virus particle.
Relations with Cells and Tissues
Many kinds of cell are infected. Particles are mainly in the cytoplasm and may
associate with all major organelles including cell walls
chloroplasts, however, may not be true virus particles
(Shalla et al., 1975
Virus-induced RNA replicase of M. Wt c. 1.6 x 105
(Ralph & Wojcik, 1969;
Zaitlin et al., 1973)
and double-stranded RNA
(Nilsson-Tillgren et al., 1974)
are associated with cytologically unspecified cell membranes.
Virus coat protein can be associated with chloroplasts
(Shalla et al., 1975),
ground cytoplasm and nuclei
(Langenberg & Schlegel, 1969;
Shalla & Amici, 1967).
Synthesis of several proteins is stimulated in infected cells
(Zaitlin & Hariharasubramanian, 1972).
Nevertheless, evidence for sites of synthesis or assembly
of the virus is equivocal.
Virus-related inclusions, visible by light and electron microscopy, range from hexagonal
crystal line plates of virus particles
to lateral aggregates of closely packed
to linear aggregates of needles, spindles and fibres, and very
large unbounded, amorphous and vacuolated X-bodies - of unknown function - composed
variously of virus particles, proteinaceous granules and tubules together with host
Type strain used in many laboratories throughout the world apparently has a common
origin. Personal recollections of C. A. Knight, W. C. Price and F. O. Holmes suggest
that the original isolate used by W. M. Stanley came from J. Johnson of the Univ. of
Wisconsin via L. O. Kunkel. The U1 strain
(Siegel & Wildman, 1954
and the German
(Wittmann-Liebold & Wittmann, 1967
also came from Johnson. TMV
was the first virus to be purified
shown to contain RNA
(Bawden & Pirie, 1937
reassembled from its constituents
(Fraenkel-Conrat & Williams, 1955
and used for production of chemically-induced mutants as a confirmation of the genetic
(Gierer & Mundry, 1958
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Leaves from healthy (upper) and systemically infected
(lower) plants of Nicotiana tabacum cv. Turkish. (Courtesy A. F. Ross.)
X-body inclusion in living leaf-hair cell of N. tabacum cv. Turkish
Samsun. Bar represents 4 µm.
Hexagonal virus crystal (left) and cell nucleus (right) in
leaf-hair cell of N. tabacum. Bar represents 6 µm.
Lateral aggregates of virus in layered ranks in parenchyma cell of N.
tabacum (courtesy K. Esau, by permission of Univ. of Wisconsin Press). Bar
represents 670 nm.
Purified virus particles negatively stained with phosphotungstate. Bar
represents 270 nm.
End view of virus fragment, negatively stained with vanadatomolybdate,
showing 3 concentric regions. Bar represents 10 nm.
Transverse section of virus particle in parenchyma cell of N. tabacum
(courtesy K. Esau, by permission of Univ. Wisconsin Press). Bar represents 11 nm.
Negatively stained virus particle showing central canal. Bar represents 48 nm.
Shadowed virus particle fragment partially degraded by phenol, showing
exposed RNA (courtesy M. K. Corbett). Bar represents 40 nm.
Model representing portion of a particle, showing helically arrayed
ellipsoidal protein subunits. Subunits in centre are removed to show arrangement of
RNA. (From work of R. E. Franklin.)