Systemic movement of Cucumber green mottle mosaic virus (CGMMV) in cucumber plants was shown to be from photoassimilate source to sink, thus indicating phloem transport. Nevertheless, CGMMV was not detected by immunocytochemical procedures in the intermediary cell-sieve element complex in inoculated cotyledons, where photoassimilate loading occurs. In stem internodes, CGMMV was first localized in the companion cells of the external phloem and subsequently in all tissues except the medulla, therefore suggesting leakage of the virus from, and reloading into, the transport phloem during systemic movement. In systemically infected sink leaves, CGMMV was simultaneously detected in the xylem and phloem. Interestingly, CGMMV accumulated to high levels in the differentiating tracheids of young leaves implying that the xylem could be involved in the systemic movement of CGMMV. This possibility was tested using plants in which cell death was induced in a portion of the stem by steam treatment. At 24 6C, steam treatment effectively prevented the systemic movement of CGMMV, even though viral RNA was detected in washes of the xylem above the steamed internode suggesting that xylem circulation occurred. At 29 6C, CGMMV systemically infected steam-treated cucumber plants, indicating that CGMMV can move systemically via the xylem. Xylem transport of CGMMV was, however, less efficient than phloem transport in terms of the time required for systemic infection and the percentage of plants infected.
INTRODUCTIONSystemic transport through the vascular system is a crucial step in plant virus infection. Most plant viruses are thought to move systemically through the phloem in parallel with photoassimilate transport (reviewed by Haywood et al., 2002;Lucas & Wolf, 1999;Nelson & Van Bel, 1998;Oparka & Turgeon, 1999;Thompson & Schulz, 1999). Phloem transport includes the loading (entry) of the virus into the phloem at source tissues, its circulation in the transport phloem and its unloading (exit) from the phloem at the sink tissues. Photoassimilate loading occurs presumably in the minor veins of source organs (Nelson & Van Bel, 1998;Santa Cruz, 1999) and in the case of cucurbits is thought to follow a symplastic route involving the specialized intermediary cell-sieve element complex (Turgeon, 1996). Knowledge of phloem circulation of plant viruses is far from complete, yet it is known that it parallels photoassimilate flow (Leisner et al., 1992;Oparka & Turgeon, 1999). The requirement of a functional capsid protein (CP) for systemic movement is common but not universal, depending on specific virus-host interactions (e.g. Dalmay et al., 1992;McGeachy & Barker, 2000;Petty & Jackson, 1990;Ryabov et al., 2001), and for most viruses the precise form in which the virus is transported has not been determined. Phloem unloading in the sink organs also parallels photoassimilate unloading and proceeds basipetally in the developing leaf during the sink-to-source transition (Cheng et al., 2000;Leisner et al., 1992; Roberts et al., 1997; Santa Cruz e...