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Garlic Mite-Borne Mosaic Virus

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Garlic Mite-Borne Mosaic Virus 日 植 病 報 62: 483-489 (1996) Ann. Phytopathol. Soc. Jpn. 62: 483-489 (1996) Characterization of a New Virus from Garlic (Allium sativum L.), Garlic Mite-borne Mosaic Virus Kazuo YAMASHITA*, Junichi SAKAI** and Kaoru HANADA** Abstract Flexuous, filamentous virus particles with a length of 700-800nm were detected in garlic (Allium sativum L.) plants showing mosaic symptoms. The virus was sap-transmissible from garlic plants to seven out of 25 plant species tested, inducing local lesions on Chenopodium morale and Gomphrena globosa and a systemic mosaic on garlic. This virus was transmitted by the eriophyid mite, Aceria tulipae Keifer, but not by aphids. Two polypeptides with MW. 30kDa and 28.5kDa were detected by SDS-PAGE of the purified virus preparation. The virus exhibited no serological relationships with leek yellow stripe, onion yellow dwarf or garlic latent viruses. No cytoplasmic cylindrical inclusions were observed in infected leaf cells of garlic or C. morale. The 3•Œ-terminal nucleotide sequence of 2518 by was determined. It contained three putative open reading frames (ORFs) for 40kDa, 28kDa (a putative coat protein, CP) and 15kDa polypeptides. The arrangement and amino acid sequences of these ORFs show close similar- ities with those of shallot virus X and garlic virus (GV)-A, -B, -C and -D. The putative CP of the mite-borne virus has 98% amino acid sequence homology with CP of GV-C, but only 60-67% identity with CPs of four other viruses, indicating the classification of the mite-borne virus and GV-C as the same virus. We propose the name garlic mite-borne mosaic virus (GMbMV) for the virus. (Received March 7, 1996; Accepted July 25, 1996) Key words: garlic, garlic mite-borne mosaic virus, mite-transmitted virus. reported12). Four different viral cDNAs derived from INTRODUCTION purified preparations of garlic plants with mixed infec- tions were designated as garlic virus (GV)-A, -B, -C, and A number of virus diseases and viruses of garlic -D27) . GV-A, -B, -C and -D were not isolated from the (Allium sativum) have been described28-30). Garlic latent infected garlic plants by single-lesion transfer, and their carlavirus (GLV)16) and a potyvirus, designated as garlic biological and serological properties are unknown. The mosaic potyvirus (GMV)1,16,24) have been reported to 3•Œ-terminal gene organization of SVX, GV-A, -B, -C and infect garlic in Japan. However, van Dijk29 reported -D are similar , but different from those of other that GMV in Japan is possibly a complex of three classified filamentous viruses, suggesting they form a distinct filamentous viruses-onion yellow dwarf potyvi- new virus group. rus (OYDV)2,6),leek yellow stripe potyvirus (LYSV)2) and We found that a mosaic disease of garlic prevalent in the garlic strain of onion mite-borne latent rymovirus Aomori Prefecture, Japan, was easily transmitted by the (OMbLV-G)30). We recently reported that LYSV isolated eriophyid mite (Aceria tulipae Keifer), and was associat- from garlic is closely related to GMV by biological and ed with flexuous filamentous virus particles33). In this serological tests34). In addition, Iwai et al.11) detected paper, we report the biological, physico-chemical, sero- OYDV from Japanese garlic by ELISA tests. However, logical, cytopathological and molecular characteristics OMbLV-G has not been detected in garlic in Japan. of the mite-borne virus isolated from garlic, which led us Recent reports based on nucleotide sequence analyses to the conclusion that the virus belongs to a new virus revealed the presence of novel filamentous viruses in group. commercial garlic in Japan27) and shallots in Russia31). The novel filamentous virus reported as shallot virus X MATERIALS AND METHODS (SVX)31) was serologically distinct from OYDV and shallot latent carlavirus (SLV), and did not produce Source of a mite-borne virus. A garlic (Allium granular inclusion bodies in infected plant tissues. The sativum cv. Fukuchi-howaito) plant showing mosaic complete nucleotide sequence of SVX has also been symptoms was collected from a field located in Aomori * Aomori Green BioCenter , Nogi, Aomori 030-01, Japan 青 森 県 グ リ ー ンバ イ オ セ ン タ ー ** Kyushu National Agricultural Experiment Station , Nishigoushi, Kikuchi-gun, Kumamoto 861-11, Japan 九 州 農 業 試 験 場 484 日本植物病理学会報 第62巻 第5号 平成8年10月 Prefecture in 1991. The virus was originally isolated by pelleted by centrifugation at 170,000•~g for 1.5hr. The eriophyid mite transmission from a diseased garlic bulb pellet was resuspended in 2ml of 0.02M BB, pH 8.0 to virus-free garlic plants. The virus was then transfer- containing 5mM EDTA. After a low speed centrifuga- red from infected garlic plants to Chenopodium morale tion (7000•~g for 10min), supernatant was collected as a by sap inoculation. After three successive single-lesion purified virus preparation. passages on C. morale, the virus was back-inoculated to Electron microscopy. Leaf extracts and purified virus-free garlic plants. virus preparations were observed after negative staining Mechanical inoculation tests. Infected garlic or with 1% phosphotungstic acid in distilled water, pH 7.0, C. morale leaf tissue was homogenized in 0.05M borate by Hitachi electron microscope H-7000. For ultra-thin buffer (BB), pH 8.5, containing 10mM EDTA, and used sectioning, tissue samples of the virus-infected garlic to inoculate 25 different species of plants. Six of rep- and C. murale plants were fixed in 2% glutaraldehyde in licates were used for each species and two different 0.1M phosphate buffer, pH 7.0 (PB), for 4hr, followed by experiments were done. Inoculated plants were post-fixation in 1% osmium tetroxide in 0.1M PB for 3 maintained in a greenhouse at 18-24•Ž and symptoms hr at 4•Ž. After a dehydration series with 50-100 were observed for six weeks after inoculation. ethanol, the tissues were embedded in Spurr's resin. Symptomless plants were back-inoculated to C. murale Thin sections were double-stained with uranyl acetate by sap inoculation. and lead citrate. Mite and aphid transmission tests. Eggs of Serology. Purified viruses (0.5mg) were eriophyid mites (A. tulipae) collected from garlic were emulsified with Freund's complete adjuvant (DIFCO transferred onto Welsh onion (A. fistulosum) seedlings. Laboratories) (1:1) and intramuscularly injected into a After hatching, their non-viruliferous first-instar nymphs rabbit three times at weekly intervals, followed by two were transferred and maintained on Welsh onion seed- intravenous injections at bi-weekly intervals. The rabbit lings. Mites at different growth stages were allowed an was bled 2 weeks after the final injection. Antisera to acquisition feeding period of 24hr on infected garlic OYDV (supplied by Dr. M. Kameya-Iwaki, Yamaguchi leaves and an inoculation access period of 24hr on University), LYSV (Dr. T. Maeda, Research Institute for virus-free garlic bulbs. Following test periods, the mites Bioresources, Okayama University), and GLV (Dr. I. were killed by dipping into the insecticide Pirimiphos- Sako, Tottori Fruit, Vegetable and Ornamental Crop methyl. Aphids, Acyrthosiphon solani and Myzus per- Experiment Station) were also used for serological tests. sicae, were starved for 2hr, and then allowed an acquisi- Serological properties were examined by leaf dip se- tion feeding period of 10-15min on infected garlic plants rology as described by Langenberg15), and by double and an inoculation access period of 1hr on virus-free immunodiffusion tests using 0.7% Difco noble agarose garlic plants. Following test periods, the aphids were gel in 0.1M PB, pH 7.0, containing 0.5% sodium dodecyl killed by spraying the insecticide DDVP. Transmission sulfate (SDS) and 0.05% sodium azide. tests with mites and aphids were done in a greenhouse SDS-PAGE and Western blotting. The molecu- using five mites or ten aphids per virus-free plant. lar weight of the coat protein (CP) from purified viruses Inoculated plants were maintained for three months to was determined by SDS-polyacrylamide gel electropho- observe symptoms. Symptomless plants were back- resis (PAGE) (12.5% slab gel)14). After electrophoresis, inoculated to C. murale by sap inoculation. the gel was stained with 0.1% Coomassie brilliant blue. Virus purification. Two to three months after After proteins were transferred onto nitrocellulose inoculation, infected fresh garlic leaves were homoge- (BIO-RAD Laboratories, Inc.), viral proteins were nized in a Waring blender for 5min with 6 volumes (v/ detected by enzyme-linked immunostaining as described w) of 0.5M BB, pH 8.5, containing 10mM EDTA and by Shirako and Ehara25) using virus-antiserum diluted 0.2%(v/v) thioglycolic acid, and then mixed with 10% with 20mM Tris-HC1 buffer (TBST), (0.15M NaCl, (v/v) carbon tetrachloride. After centrifugation of the 0.05% NaN3, pH 7.5, and 0.02% Tween 20) at 1: 4000 and homogenate at 7000•~g for 10min, 1% (v/v) Triton alkaline phosphatase-conjugated anti-rabbit goat im- X-100 , 5%(w/v) polyethylene glycol (mol. wt. 6000) and munoglobulin G (Sigma Chemical Company) diluted 0.1M NaCI was added to the aqueous phase. After with TBST at 1: 2000. stirring for 1hr at 4•Ž, the suspension was centrifuged at RNA extraction and cDNA cloning. Viral 7000•~g for 10min. The pellet was resuspended in RNA was extracted from purified viruses using one-fifth the original volume of the extract in a solution proteinase K and SDS followed by phenol extraction and of 0.05M BB, pH 8.3, 0.5M urea and 10mM EDTA ethanol precipitation. The molecular weight of viral (suspending buffer), then stirred gently for 1hr. After RNA was estimated by 0.8% agarose gel electrophoresis two cycles of differential centrifugation (7000•~g for 10 under denaturing conditions with 2.2M formaldehyde.
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