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Rice stripe virus disease (RSVD) occurs in the temperate regions of East Asia, specifically China, Japan, Korea, and Taiwan, and it has also been reported from far-eastern Russia (Ou 1985).
RSVD is one of the most serious diseases of rice in the temperate regions of East Asia. It can cause high yield losses when severe epidemics occur. It has affected several thousand hectares of rice-growing areas (Table 1). Severe infection at the seedling to early tillering stage was reported to cause yield losses of 50% to 100% (Lin et al 1990). In eastern China, RSVD caused yield losses of 30% to 40% in 2003-04 (Zhang et al 2007).
Table 1. Some reported epidemics of RSVD.
|Year||Country||Area affected (ha)||Reference|
|1960s||Japan||500,000||Maeda et al (2006)|
|1960s||Eastern and southern China||2,660,000||Wang et al (2008)|
|1963-67||Japan||500,000 to 620,000 (annually)||Nemoto et al (1994)|
|1986||Japan||170,000||Nemoto et al (1994)|
|Not indicated||Yunnan Province, China||67,000||Wang et al (2008)|
|2002||Jiangsu Province, China||780,000||Wei et al (2009)|
|2003||Jiangsu Province, China||957,000||Wei et al (2009)|
|2004||Jiangsu Province, China||1,571,000||Wei et al (2009)|
|2005-06||Zhejiang Province, China||100,000||Wang et al (2008)|
|2007-08||Korea||84% of rice fields||Jonson et al (2009)|
|2007||Zhejiang Province, China||17,600||Zhu et al (2009)|
RSVD causes chlorotic to yellowish white stripes, mottling, and necrotic streaks on the leaves (Fig. 1). Plants that are infected at the seedling stage have folded, twisted, wilted, and droopy leaves; are stunted; have few tillers; may produce few panicles; and may die prematurely (Webster and Gunnell 1992). Panicles produced by infected plants have whitish to brown and deformed and unfilled spikelets, and may not be fully exserted. Leaves of plants that are infected at the maximum tillering stage or older have less severe chlorosis or mottling. Panicle exsertion and ripening of these plants may be delayed.
Rice stripe virus (RSV) is the type member of the genus Tenuivirus. It is filamentous or thread-like, 500–2,000 nm in length, with a diameter of only 3 to 8 nm. The virion consists of four segmented nucleocapsids. These single-stranded RNA segments are the following:
RNA 1—8.9 kb, completely negative sense, contains a single open reading frame and encodes RNA-dependent RNA polymerase (Toriyama et al 1994), which is known to be responsible for replication and transcription of the viral RNA genome.
RNA 2—3.5 kb, ambisense, encodes two nonstructural proteins: a membrane-associated protein in the viral or positive sense and a putative membrane glycoprotein in the viral complementary or negative sense (Takahashi et al 1993).
RNA3—2.7 kb, encodes a nonstructural protein (NS3) that functions as a suppressor of gene silencing in the viral sense and the nucleocapsid protein in the viral complementary sense (Kakutani et al 1991, Xiong et al 2008).
RNA4—2.1 kb, encodes two nonstructural proteins: a protein known as a major noncapsid protein in the viral sense that accumulates in infected plants and may be involved in pathogenesis and a nonstructural protein that functions as a movement protein in the viral complementary sense (Toriyama 1986, Kakutani et al 1990, Zhu et al 1992, Xiong et al 2008).
RSV does not produce enveloped virions in infected plants (Liang et al 2005).
The virus is transmitted in a persistent, circulative-propagative manner mainly by the small brown planthopper (SBPH), Laodelphax striatellus Fallén. It is also transmitted by three other planthopper species, Unkanodes sapporona (Matsumura), U. albifascia (Matsumura), and Terthron albovittatum (Matsumura). The virus is transmitted from female adults to offspring through the eggs. It can be transmitted to about 90% of the offspring for up to 40 successive generations (Lee 1969). The virus can be transmitted by mechanical inoculation, but with difficulty. It is not transmitted by seeds or by contact between plant tissues.
The optimum acquisition period is 1 day, but the acquisition period can be as short as 15 minutes. The incubation period of the virus in L. striatellus is usually 5 to 10 days; however, it can be as long as 21 days. Females transmit the virus more efficiently than males. The ability of L. striatellus to transmit the virus decreases with age (Iida and Shinkai 1969) and after overwintering. However, overwintered insects maintain a high transmission rate to their progeny (Iida and Shinkai 1969).
The optimum temperature for transmission of the virus is 25–30°C (Chung 1974).
The virus multiplies in the vector and is retained when it molts.
RSV infects around 80 species of the family Gramineae and several nongraminaceous species, including wheat, barley, foxtail millet, rye, and oat (Ou 1985, Hibino 1996). RSVD epidemics in some areas have been attributed to the increase in area planted to wheat and barley. Cereals, especially wheat and barley, are not considered as important reservoirs of the virus but they serve as habitats for the vector during the rice fallow period (Hibino 1996). After rice is harvested, some of the viruliferous adult vectors migrate to grasses surrounding rice fields or nearby fields and oviposit. Congenitally infected nymphs migrate to newly established crops and to weeds and overwinter as fourth-instar nymphs and diapause (Hibino 1996). First-generation adults of this overwintering generation emerge and some move to newly established rice fields. Second-generation adults appear about a month later. First- and second-generation adults feed on rice at seedling and early tillering stages, respectively, and may cause severe epidemics (Wang et al 2008). On the other hand, the viruliferous vectors can transmit the disease from rice to newly established seedlings of wheat and barley in winter, causing severe epidemics in these crops as well (Xiong et al 2007).
Japonica varieties that are grown in lowland areas are generally susceptible to RSVD, whereas indica, Javanese, and japonica upland varieties have genetic resistance. These resistant upland varieties have genes with dominant resistance to RSV, Stv-a, and Stv-b (Washio et al 1968). Indica varieties have Stvb-i, which is allelic with Stv-b. Varieties that are currently grown by farmers have only the Stvb-i gene (Nemoto et al 1994), which has remained resistant to RSV since the 1960s (Maeda et al 2006).
QTLs that confer resistance to RSV (Wang et al 2011, Zhang et al 2011, Wu et al 2011) and tolerance to the vector (Zhang et al 2010) have been identified. Some of these QTLs could be pyramided through marker-assisted selection.
Growing resistant varieties is the most economical and practical approach in managing RSVD. Resistance to the virus is more effective in controlling RSVD than resistance to the vector (Okamoto and Inoue 1967). Crop establishment should be timed so that the crop will be at the stem elongation stage or older during the peak of immigration of viruliferous insects from winter crops, specifically wheat and barley, as plants at the seedling to early tillering stages are highly susceptible to RSVD (Wang et al 2008, Zhu et al 2009).
Insecticides should be applied judiciously to reduce the population of viruliferous vectors. Indiscriminate application of insecticides has resulted in the resistance of populations to certain compounds (Otuka et al 2010). Synchronous planting should be practiced over wide areas. RSVD epidemics in Japan in the 1960s were partly attributed to staggered planting of rice (Hibino 1996). Ratoon or stubbles of the previous crop and weeds should be removed to reduce the virus and the population of the vector.
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Developed with input from: NP Castilla, S Savary, A Sparks, and I-R Choi.