Hordeivirus relationships and genome organization

The hordeiviruses are a unique group of RNA viruses composed of three members (25, 62, 75): barley stripe mosaic virus (BSMV), poa semilatent virus (PSLV), and lychnis ringspot virus (LRSV); with a possible fourth member, anthoxanthum latent blanching virus (ALBV). The short rigid virus rods of hordeiviruses are larger in diameter and about half the length of tobacco mosaic virus, but they have a much greater range of variation in their length. The rods are composed of protein subunits encapsidating single­ stranded RNAs of variable size that comprise about 4% of the virion. Although members of the group have similar particle structure (40), they exhibit considerable biological diversity (25, 62, 79). BSMV (79), PSL V (97) and ALBV (28) primarily infect members of the Gramineae, while LRSV infects plants in several dicot families (13). Vector relationships have not been established for any of the hordeiviruses, and carry-over of BSMV from year to year depends primarily on seed transmission. Although many field isolates of BSMV are efficiently seed transmitted, other strains are in­ frequently transmitted through seed when tested in the laboratory (23, 101). LRSV is highly transmissible through both seed and pollen (14), but seed transmission has not been demonstrated for either PSL V (97) or ALB V (28). Our main objective in this review is to discuss the relationships among the hordeiviruses and to describe the structural properties of the BSMV genome that distinguish it from other tripartite RNA viruses . The biology and patholo-

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