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Presence of Brome mosaic virus in Barley Guttation Fluid and Its Association with Localized Cell Death Response

Water exits from inside the leaf through transpiration or guttation. Under conditions to promote guttation, surface fluid (guttation fluid) from Brome mosaic virus (BMV)-infected barley, wheat, and maize plants was analyzed for the presence of the virus by biological and serological assays. We also investigated the route by which BMV exited infected cells to the intercellular space of the barley leaf. BMV was detected in guttation fluid from systemically infected barley leaves when the initial viral symptoms were observed on these leaves. The virus was also detected in guttation fluid from systemically infected wheat leaves, but not in maize leaves showing either systemic necrosis or chlorotic streaks. Interestingly, in BMV-infected barley leaves, but not in maize leaves showing chlorotic streaks, cell death occurred within and adjacent to veins. Staining of xylem and phloem networks in infected barley leaves with fluorescent dyes showed that xylem, and to a lesser extent phloem, were severely damaged and thus became leaky for dye transport. No such damage was observed in BMV-infected maize leaves showing chlorotic streaks. We propose that in infected barley leaves, BMV exits from damaged vein cells (especially the xylem elements), accumulates in intercellular spaces, and then reaches the surface of the leaves through stomata during guttation or transpiration. In nature, BMV may be carried to adjacent plants and cause infection by movement of vertebrate and invertebrate vectors among infected plants exuding guttation fluid.