Complete sequence and genetic characterization of Raspberry latent virus, a novel member of the family Reoviridae☆,☆☆
Introduction
The family Reoviridae contains viruses with genomes composed of multiple (9–12) segments of linear double-stranded RNA (dsRNA). This family is the most diverse of the dsRNA virus families, as it includes species isolated from a wide range of hosts including mammals, birds, fish, insects, arachnids, marine protists, crustaceans, fungi and plants (Brussaard et al., 2004, Hillman et al., 2004, Mertens et al., 2005).
More than 75 virus species have been classified as members of 12 well-established genera: Orbivirus, Orthoreovirus, Rotavirus, Coltivirus, Aquareovirus, Cypovirus, Fijivirus, Phytoreovirus, Oryzavirus, Seadornavirus, Mycoreovirus and Idnoreovirus (Mertens et al., 2005). The genera Cardoreovirus, Dinovernavirus, and Mimoreovirus, have been established recently and include viruses isolated from crabs, mosquitoes and marine protists, respectively (Attoui et al., 2005, Attoui et al., 2006, Zhang et al., 2004).
Currently, there are three genera of plant reoviruses; Fijivirus, Oryzavirus and Phytoreovirus. The genus Fijivirus includes members with 10 segments and is divided into 5 groups based on host, vectors and serological properties (Isogai et al., 1998a, Mertens et al., 2000). Members include viruses that infect monocots and insects (Hunter et al., 2009, Nakashima et al., 1996, Noda and Nakashima, 1995). The genus Oryzavirus includes two members; Rice ragged stunt virus (RRSV) and Echinochloa ragged stunt virus (ERSV), also comprised of 10 segments (Chen et al., 1989, Holmes et al., 1995, Shikata et al., 1979, Uyeda and Milne, 1995, Yan et al., 1992). Phytoreovirus includes viruses with 12 segments that infect both monocots and dicots with Wound tumor virus (WTV), Rice dwarf virus (RDV) and Rice gall dwarf virus (RGDV) being the recognized members of the genus (Black, 1945, Mertens et al., 2000, Nuss and Dall, 1990). The recent characterization of Tobacco leaf enation virus (TLEV) (Picton et al., 2007, Rey et al., 1999), and Homalodisca vitripennis reovirus (HoVRV), isolated from the hemipteran Homalodisca vitripennis (Stenger et al., 2009), have expanded this genus now to include insect viruses.
In 1988, a novel reovirus was identified in red raspberry in Washington State, USA. Partial sequence was obtained and used to design detection primers (Jelkmann and Martin, 1989). The virus has been detected commonly in the main raspberry production areas in the Pacific Northwest (PNW), especially in areas where raspberry crumbly fruit disease is prominent (Murant et al., 1974, Martin, 1999).
The new reovirus does not cause symptoms when grafted onto raspberry indicator plants and thus, the name Raspberry latent virus (RpLV) is proposed. The characterization of RpLV has been necessary in order to elucidate possible interactions with other viruses and its implication in crumbly fruit and other raspberry diseases.
Section snippets
Type isolate source
Root cuttings from ‘Meeker’ raspberry were obtained from production fields in northern Washington State, USA. The cuttings were originated from 3-and 4-year-old plants, some of which had shown crumbly fruit symptoms in previous seasons. A total of 90 root cuttings were planted, grown in 1 gal pots and maintained in a greenhouse under standard conditions (12 h day light at 25 °C). It is important to point out that we did not know the status of each cutting at the time of planting, i.e. whether each
Electrophoretic analysis of double-stranded RNAs
Double-stranded RNA from RpLV singly-infected plants was separated by 1.5% agarose gel electrophoresis (AGE) and 10% PAGE. The AGE profile was similar to those of other reoviruses and revealed the presence of multiple segments arranged in three groups, based on their molecular masses (high, intermediate and low) (Fig. 1). The PAGE profile allowed for the separation of three distinct dsRNA segments that co-migrated as a single band in AGE. S1–S4 form the high molecular mass group, the
Discussion
RpLV was first isolated from raspberry in 1988 and has been found at high incidence in the main production areas in the northern Washington State, USA and British Columbia, Canada. The suggested name derives from the fact that RpLV is symptomless in single infections in standard Rubus virus indicators or in the red raspberry cultivars tested. However, RpLV is implicated in a new virus complex responsible for causing severe crumbly fruit in red raspberries (Quito-Avila et al., 2009). The high
Acknowledgements
This work was funded partially by the US Department of Agriculture under the Specialty Crops Research Initiative (SCRI), award number 2009-51181-06022. We are grateful to Dr. Sead Sabanadzovic from Mississippi State University for sharing unpublished data on a closely related reovirus, Simone Alt and Svenja Leible (Julius Kuhn Institute for plant protection in fruit crops and viticulture, Dossenheim, Germany) for initial sequencing work, and to growers in northern Washington State for providing
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