Rice stripe virus
Department of Microbiology, National Institute of Agro-Environmental Sciences, Kannondai 3, Tsukuba, Ibaraki 305-8604, Japan.
Disease described by
Virus characterized by
S. Toriyama (1982a
S. Toriyama et al. (1994a)
A virus with filamentous particles, 500-2000 nm long, but only about 8 nm wide. The particles sediment as
four components and contain four ssRNA species, a major protein (the nucleocapsid protein) and a minor protein
(the RNA polymerase). RNA-1 is negative-stranded and RNAs 2 to 4 are ambisense. Non-structural protein material
is produced abundantly in infected cells. Mechanical inoculation is difficult. The virus is transmitted
by Laodelphax striatellus and three other planthopper species in a persistent manner; it is transmitted
through the egg to about 90% of progeny insects. It infects many species of Gramineae. It occurs in rice-growing
areas of Asia and the USSR, and causes significant reduction in rice yield.
In rice the virus causes chlorotic stripes, chlorosis, moderate stunting and loss of vigour
in severe infections the leaves develop brown to grey necrotic streaks and die. Diseased plants produce
few or no panicles: those produced carry whitish to brown discoloured malformed spikelets
). Early infection of rice causes significant loss of yield
(Yasuo, Ishii & Yamaguchi, 1965
late infection also reduces yield by retarding ear emergence and ripening
(Yasuo et al., 1965
In maize and wheat the virus causes chlorotic stripes, chlorosis and stunting.
The disease was first recognized in the early 1900s in central Japan, where severe damage was caused to
rice crops. Since the 1950s the extension of early planting favoured the occurrence of the disease in
(Yasuo et al., 1965
and during the 1960s and 1970s about 200,000 ha of rice was affected each year. In Korea, the disease
affected 40% of rice hills in 1965
Stripe disease incidence in Taiwan was most severe around 1970: 1045 ha were affected in 1973
Stripe disease is broadly prevalent all over the rice-growing areas in China
Lin et al., 1990
and the USSR
(Reifman, Pinsker & Krylova, 1978
Host Range and Symptomatology
RSV occurs naturally in rice, maize, wheat, oat, foxtail millet and wild grasses such as
Cynodon dactylon, Digitaria adscendens, D. violascens, Eragrostis multicaulis
The virus is reported to infect 37 species of the family Gramineae.
The main type of symptom induced is chlorotic striping
Yamada & Yamamoto, 1956
Non-gramineous plant species, such as Cyperus amuricus
, C. sanguinolentus
and Eriocarlan robustius
are also reported as host plants
(Chung & Lee, 1971
Pinsker & Reifman, 1975
The virus is readily transmitted to test plants by the planthopper, Laodelphax striatellus,
is difficult to transmit mechanically.
Okuyama & Asuyama (1959)
obtained infection in 6% of plants that were injected with crude extracts made from diseased plants
with 0.01 M cysteine-HCl, and
achieved infection in 0.9% of plants that were injected with extracts of viruliferous insects.
Oryza sativa (rice). Seedlings of most Japanese paddy varieties are highly susceptible to infection.
Chlorotic stripes, mostly light yellowish broken streaks
(Fig.1), develop on systemically infected leaves 10 to 25 days after inoculation
by planthoppers (L. striatellus). Characteristically, limp chlorotic leaves emerge without
unfolding, elongate, droop and wilt
(Fig.3). Yellowing and moderate stunting also occur.
Triticum aestivum (wheat) and Zea mays (maize) (cv. Golden Cross Bantam).
Symptoms in these species are similar to those in O. sativa. Wheat plants may produce
whitish, rolled, needle-like leaves that quickly droop.
Rice, wheat and maize are used to maintain cultures and as a source of virus for purification.
Young rice seedlings are readily killed, and so care must be taken to use the appropriate growth stage
and varieties, e. g. third leaf stage of rice cv. Norin No. 8 or Kinmaze. When maize seedlings
are used, wheat, barley or rice plants must also be planted, or the planthoppers will die
within 2-3 days.
Young rice and wheat seedlings are suitable for assaying transmission by insect vectors.
An isolate that causes mild symptoms on rice
(Ishii & Ono, 1966
and two variants that differ in symptoms induced in wheat and in insect transmissibility
are reported. Also,
Hayashi et al. (1989)
found four isolates that produce disease-specific non-structural proteins of different molecular weights.
However, there are no detailed studies of any of these isolates.
Transmission by Vectors
The most active vector in the field is the small brown planthopper, Laodelphax striatellus
three other planthopper species, Unkanodes sapporona
and Terthron albovittatum
also transmit. The proportion of active transmitters of L. striatellus
is about 20%
Although the shortest acquisition feeding period is 15 min, best transmission is obtained with
planthoppers that acquire the virus by feeding for 1 day. The incubation period of the virus in
ranges from 5 to 21 days but incubation is complete within 5 to 10 days for
most individuals. Minimum inoculation feeding time is 3 min: about half or more of the infective
planthoppers infect rice seedlings after feeding for 1 h
(Yamada &Yamamoto, 1955
Ability of the insects to transmit the virus decreases markedly with age. Females of
are more efficient transmitters than males
The virus passes through a high percentage of eggs to progeny, about 90% in
Yamada &Yamamoto, 1954
was selected and bred for high or low ability to acquire and transmit the
virus (50-60% and less than 10% of the insects transmitting, respectively), and this was correlated
with the frequency of transmission of the virus through the eggs
Evidence of virus multiplication in L. striatellus
has been obtained by repeated passage through
the eggs for 6 years through 40 generations
by serial transfer of the virus from insects to insects by injection
(Okuyama, Yora & Asuyama, 1968
and by detection of virus particle antigen in various organs of viruliferous insects with the
fluorescent antibody staining technique
(Kitani, Kiso & Yamamoto, 1968
Transmission through Seed
None found in rice
RSV is a good immunogen. Rabbit antisera with titres of 1/512 to 1/1024 in precipitin tests are
readily obtained. In agar gel diffusion tests, it is necessary to disrupt virus particles in
purified preparations or crude sap of infected tissues by treatment with 0.5% SDS. The haemagglutination
test was used to detect the virus particle antigen at high dilutions in vectors or plants
(Saito & Iwata, 1964
Yasuo &Yanagita, 1963
Sonku & Sakurai, 1973b
Monoclonal antibodies have been made and used for specific detection of the viral antigen
(Omura et al., 1986
and ELISA is useful for detecting the virus in plants and vectors
(Omura et al., 1986
Y. Takahashi et al., 1987
Cheng et al., 1991
Xie et al., 1991
For practical applications, the latex flocculation test
(Omura et al., 1984
rapid immunofilter paper assay
(Cabauatan et al., 1994
and dot immunobinding assay (DIBA)
(Qin, Gao & Cheng, 1994
) are useful.
RSV is the type member of the genus Tenuivirus
. Serologically, it is moderately closely related
to maize stripe virus
(Gingery, Nault & Yamashita, 1983
and distantly related to
rice grassy stunt virus
(Hibino et al., 1985
but not related to
rice hoja blanca virus
(Yasuo, Yanagita & Yamaguchi, 1961
The amino acid sequences encoded by ORFs on RNAs 2 to 4 had a moderately high similarity to those encoded
by the corresponding ORFs of maize stripe and rice hoja blanca viruses, but very low similarity to
those encoded by the ORFs on RNAs 2, 5 and 6 of rice grassy stunt virus
(S. Toriyama et al., 1994a
RSV has no ORF corresponding to the ORFs on RNAs 3 and 4 of RGSV. On the other hand, the amino acid
sequences of the ORFs encoded on RSV RNA-1 and RGSV RNA-1 are highly conserved. The 5' and 3' termini of
RSV RNAs 1-4 contain 8-nucleotide complementary sequences which are identical to those of animal viruses in
the genus Phlebovirus
(M. Takahashi et al., 1990
and the amino acid sequence of the RSV RNA polymerase protein has appreciable homology (about 30% identity
over 1500 residues) with the polymerase protein of phleboviruses
(S. Toriyama et al., 1994a
Stability in Sap
Assayed by observing transmission by injected insects. Dilution end point:
in extracts from viruliferous insects and
in sap from diseased leaves. Thermal inactivation point (5 min):
50-55 șC. Longevity: 4 days in extracts of viruliferous insects kept at 4 șC, 8-12 months in
viruliferous insects and diseased rice plants kept at -20 șC, and 1-2 months in purified preparations
kept at -20 șC
(Kiso, Yamamoto & Kitani, 1974
(S. Toriyama, 1982a
Grind infected leaves in 0.1 M Na2
containing 10 mM DIECA, and adjust to pH 7.2
with solid ascorbic acid. After clarification by treatment with 20% chloroform, collect the virus
by centrifugation for 2 h at 123,000g
or by precipitation from 8% polyethylene glycol.
Resuspend the virus in 0.01 M phosphate buffer, pH 7.5, and purify by repeated low and high
speed centrifugation. Host plant impurities can be precipitated by adding solid ammonium sulphate to
30% saturation. The preparation can be further purified by centrifugation in 10-40% linear sucrose
density gradients. The infective component (nB) aggregates readily and sediments to the bottom of
sucrose gradient tubes: it may be necessary to resuspend the pellet and repeat the density
gradient centrifugation. The yield of virus is 20-30 mg per 100g fresh leaves. Addition of
, EDTA or bentonite to the buffer did not improve preservation of the
virus particles (S. Toriyama, unpublished data).
Properties of Particles
The virus particles sediment as three main components in sucrose density gradients: middle (M), bottom (B)
and nB (Fig.4
(S. Toriyama, 1982a
However, M component can be partially separated into two components, M1 and M2
(Ishikawa, Omura & Tsuchizaki, 1989b
A 'top component' also occurs but consists of degraded particles
(S. Toriyama, 1982a
Sedimentation coefficients (s20,w), determined on a preparation with
A260=10: 72 S (M)
65 S (M), 80 S (B) and 98 S (nB) (S. Toriyama, unpublished data).
A260/A280: 1.49, determined on an unfractionated preparation
(Koganezawa, Doi & Yora, 1975).
Absorbance at 260 nm (1 mg/ml, 1 cm light path): 4.4, determined on an unfractionated preparation;
not corrected for light-scattering (S. Toriyama, unpublished data).
Buoyant density in caesium chloride: 1.282 g/cm3. This single component contains all
the components that separate in sucrose density gradients (S.Toriyama, unpublished data).
Isoelectric point: particles precipitate at around pH 4.5 (S. Toriyama, unpublished data).
The virus was formerly thought to have spherical particles
(Koganezawa et al., 1975
but it is now known that the particles are basically filamentous, although they are frequently observed
as apparently branched and other pleomorphic structures
(Koganezawa et al., 1975
S. Toriyama, 1982b
The modal length of circular filaments found in the MI, M2, B and nB centrifugal components was 510, 610,
840 and 2110 nm, respectively
(Ishikawa, Omura & Hibino, 1989a
The viral structure is extremely fragile and readily unfolded. The branched appearance is caused
by supercoiling: completely unfolded particles possess a helical structure 3 nm wide which in turn
forms secondary coils about 8 nm wide
(S. Toriyama, 1982b
These unfolded pleomorphic particles folded to form stiff rod-like particles, 8 nm wide and/or
circular filaments under high salt conditions
(S. Toriyama et al., 1994b
Particle CompositionNucleic acid
(S. Toriyama, 1982a
However, dsRNAs containing the sequence of viral ssRNAs are also detected in agarose gel electrophoresis
(S. Toriyama & Watanabe, 1989
Ishikawa et al., 1989b
The two smallest RNAs, RNA-3 and RNA-4, occur in M component (which, as mentioned above, is a mixture of
two components, M1 and M2), and RNA-2 occurs in B component. The nB component contains RNAs 2 to 4
together with the largest RNA, RNA-1, and only this nB component is infectious (Fig.8
(S. Toriyama, 1982a
The RNA constitutes about 12% of the particle weight, as judged by the
value of purified preparations. The three RNA species from M and B
components are linear molecules about 0.7-1.0 ”
(S. Toriyama, 1982a
Protein: One species of nucleocapsid protein, of 32 KDa
S. Toriyama, 1982a).
The sequence of 322 amino acid residues is available in the database accession numbers,
(Kakutani et al., 1991;
Zhu et al., 1991).
A minor, but large molecular weight protein, RNA-dependent RNA polymerase, is also associated with
(S. Toriyama, 1986),
and the amino acid sequence is available (database accession number D31879)
(S. Toriyama et al., 1994).
Other components: No significant amounts of lipid or polyamine are found in purified
preparations (S. Toriyama, unpublished data).
Database accession numbers of the genome: RNA-1,
(China). The sequences of about 20 bases at the 5'-
and 3'-termini of each RNA are almost complementary to each other: ten nucleotides of these termini
are identical among RNAs 1 to 4, except for one base change in RNA-1
(M. Takahashi et al., 1990
The genome organization is shown in Fig.9. RNA-1, which encodes the
RNA-dependent RNA polymerase, is negative-stranded
(S. Toriyama et al., 1994a)
and RNAs 2 to 4 are ambisense
(Kakutani et al., 1990,
Zhu et al., 1991,
M. Takahashi et al., 1993).
In each of RNAs 2 to 4, the intergenic non-coding region between the two ORFs contains an oligo(A)-
and oligo(U)-rich sequence that can be formed into base-paired hairpin stems
(Zhu et al., 1991,
The nucleocapsid protein is encoded at the 5'-proximal region of virus complementary sense RNA-3, and the
major non-structural protein is encoded at the 5'-proximal region of virus sense RNA-3. The major
non-structural protein is synthesized in an in vitro translation system by using the viral RNA,
but nucleocapsid protein is not synthesized
(Hamamatsu et al., 1993).
mRNAs with the genomic size of these proteins are produced in vivo, and they have an additional 10
to 23 non-viral nucleotides at the 5'-termini
(Shimizu et al., 1996).
At present there are no experimental data on the functions of the ORFs on viral sense RNAs 2 & 3, or
on viral complementary sense RNA-4.
Relations with Cells and Tissues
Large inclusions, shaped like rings, rods or figures of eight, are present in infected cells
Hirai et al., 1964
Reifman et al., 1978
The inclusions usually contain many granules, but some have no granules and resemble crystalline
inclusions. The inclusions probably consist of the non-structural protein which is serologically
unrelated to the coat protein and is produced abundantly in infected cells of plants with severe
symptoms but less so or not at all in infected cells of tolerant or resistant varieties. The isoelectric
point of purified non-structural protein is pH 5.4. The protein forms needle crystals at pH 4-5.
Its sedimentation coefficient (s20,w
) is 3
(Kiso & Yamamoto, 1973
One polypeptide species, of 20 KDa, was reported by
The sequence of 286 amino acid residues is available in the database accession numbers
(Kakutani et al., 1990
Zhu et al., 1992
In sections of infected plant cells
recognition of individual filamentous virus particles is difficult but granular regions, sometimes enclosed
by membranes, can be observed in the cytoplasm. These regions are possibly formed from virus
aggregates. Virus particle antigen was detected in phloem tissue and mesophyll of infected wheat leaves
by fluorescent antibody staining
(Kiso et al., 1974).
The virus moves downwards in the phloem at a rate of 25-30 cm/h at 30 șC and multiplies when young
tissues are reached
(Sonku & Sakurai, 1973a,
In viruliferous insect cells, examined by immunogold labelling of thin sections, the viral antigen
was detected on amorphous or filamentous regions in the cytoplasm of the midgut, the principal
salivary gland, the ovarioles and the fat body
(Suzuki et al., 1992).
Ecology and Control
An epidemic of rice stripe disease in Japan was triggered by the wide introduction of the earlier
cultivation of rice
(Yasuo et al., 1965
It is likely that the earlier transplantation of rice seedlings coincided with the migration date of the
vector L. striatellus
, which had earlier multiplied in wheat fields. Thus, transplanting or
seeding date affects disease incidence
(Yasuo et al., 1965
Bae & Kim, 1994
The resistance of rice (Oryza sativa) to RSV has been extensively studied. Most Japanese
paddy varieties are highly susceptible but Japanese upland varieties and Indica-type rice varieties
are resistant and/or tolerant. The resistance is directed against the virus, not against the insect
vector. Highly resistant hybrids, Chugoku No. 31 and St No. 1, were bred by crossing cv Norin
No. 8 (Japonica) and cv Modan (Indica), and then back-crossing 8 times with cv Norin
No. 8. These hybrids have been used as mother lines for breeding practical new resistant
varieties. Inheritance of resistance appears to be governed by multiple alleles with various levels
of resistance: gene St2 in Japanese upland varieties and gene St2i
in Indica-type varieties. A gene St1 at another locus as well as
St2 is essential for resistance in Japanese upland varieties
(Yamaguchi, Yasuo & Ishii, 1965;
Sakurai & Ezuka, 1964;
Washio et al., 1967,
K. Toriyama, 1969,
K. Toriyama et al., 1966,
The cultivation of resistant varieties is favoured for decreasing the density of viruliferous insects
(Takayama et al., 1987).
RSV is readily distinguished from other viruses infecting rice, wheat and maize by its characteristic
symptoms and by its transmissibility by L. striatellus.
The virus, together with other tenuiviruses, is evolutionarily related to phleboviruses of the
(S. Toriyama et al., 1994a).
Viruses in both genera have multipartite genomes possessing negative sense and antisense components.
The 5' and 3' ends of the genomic RNAs have complementary sequences of which eight nucleotides are identical
in the two genera. Like viruses in most genera of Bunyaviridae, tenuiviruses infect their
insect vectors as well as their primary hosts. The number of genome components and the apparent absence
of a membrane-bound particle distinguish tenuiviruses from viruses in the family Bunyaviridae.
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Chlorotic stripe symptoms on infected leaves of rice (Oryza sativa) compared with a healthy leaf (extreme left).
Malformed and immature ears, and chlorotic stripes on flag-leaves, of rice plants infected at a late growth stage.
Drooping and wilting symptoms in young seedlings of O. sativa (cv. Norin No. 8).
Bands formed by rice stripe virus after sedimentation in a sucrose gradient column: middle component (M), bottom component (B) and nB component (nB).
Filamentous particles, c. 8 nm wide, seen in purified nB component. Bar represents 100 nm.
Partially and completely unfolded virus particles. Bar represents 100 nm.
Completely unfolded particles c. 3 nm wide. Note that they still retain a coiled appearance. Bar represents 100 nm.
Ribonucleic acid species on polyacrylamide gels: (left), three RNA segments from M+B components;
(right), four RNA segments in a preparation from nB component, all except the uppermost being
caused by contamination with M+B components.
Genome organization of RSV (isolate T). The ORFs of four RNA segments are displayed on viral-sense RNAs
as a black bar and on viral complementary- sense RNAs as a blue bar. The proteins deduced from each ORF
are shown as boxes, indicating the molecular sizes of the predicted proteins. Pol, RNA polymerase;
N, nucleocapsid protein; NSmajor, a major non-structural protein.