Pea seed-borne mosaic virus seed transmission exploits novel symplastic pathways to infect the pea embryo and is, in part, dependent upon chance

Seed transmission of pea seed-borne mosaic virus (PSbMV) depends upon symplastic transport of the virus from infected maternal cells to the embryo. Such transport pathways have not been identified in higher plants. To identify these pathways, we have studied the ultrastructure of the tissues and cells, around the micropyle of young developing seeds and compared transmitted and nontransmitted virus isolates. A characteristic of PSbMV infection was the presence of cylindrical inclusions positioned over plasmodesmal openings. The presence of cylindrical inclusions on the testa-endosperm boundary wall, together with immunogold labelling for virus-specific products on the wall and in the endosperm, indicated that symplastic connections existed at this interface. Close examination of the endosperm-suspensor boundary at the base of the suspensor revealed discontinuities in the suspensor sheath wall as porelike structures, which the virus might pass through en route to the embryo. A nontransmitted PSbMV isolate was able to invade the maternal tissues of the developing seed but was excluded from the embryo, although it was detected at a low level in the endosperm. Since the endosperm did not support virus replication, it appeared that passive accumulation determined the amount, timing, and location of the virus relative to the base of the suspensor. Rarely, therefore, could the nontransmitted virus isolate reach the correct location in the endosperm at the correct time for embryo infection via the suspensor to occur.