The genomes of umbraviruses differ from those of most other viruses in that they do not encode a coat protein, and thus no virus particles are formed in infected plants. Protection of umbraviral RNA outside the host plant, during vector transmission, utilizes the coat protein of an assistor luteovirus, but this review focuses on the mechanisms that compensate for the lack of a coat protein in processes within the host plant. As well as an RNA-dependent RNA polymerase, umbravirus genomes encode two other proteins from almost completely overlapping open reading frames. One of these is a cell-to-cell movement protein that can mediate the transport of homologous and heterologous viral RNAs through plasmodesmata without the participation of a coat protein. The other, the ORF3 protein, binds to viral RNA to form filamentous ribonucleoprotein particles that have elements of helical structure. It serves to stabilize the RNA and facilitates its transport through the vascular system of the plant. It may also be involved in protection of the viral RNA from the plant's defensive RNA-silencing response, although it is not a suppressor of silencing. The ORF3 protein also enters the cell nucleus, specifically targeting the nucleolus. Although the function of this localization is unknown, the ORF3 protein may provide a valuable tool for investigating plant nucleolar function.
INTRODUCTIONOne of the main characteristics of most viruses is the formation of virus particles or virions, in which the viral genomic nucleic acid (RNA or DNA) is protected by encapsidation with one or more types of capsid (or coat) protein (CP). In virions, molecules of CP are packed into regular uniform structures based on either helical or icosahedral symmetry. In plant viruses, CP is also involved in transmission by biological vectors and often in the spread of viruses in infected plants. Some plant viruses, such as Cowpea mosaic virus (CPMV; Wellink & van Kammen, 1989), Potato virus X (PVX; Santa Cruz et al., 1998) or Cucumber mosaic virus (CMV; Canto et al., 1997), require CP for movement both through plasmodesmata (cell-to-cell movement) and via the phloem (long-distance movement). Others, such as Tobacco mosaic virus (TMV), require CP for long-distance movement but not for cell-to-cell movement (Carrington et al., 1996). Umbraviruses are distinguished from most other viruses by their lack of a gene for CP, and as a result these viruses do not form conventional virus particles.The name of the genus Umbravirus is derived from the Latin umbra, which means shadow, both in the physical sense and in the metaphorical senses of a phantom or an uninvited guest who comes with an invited one. This name reflects the way in which umbraviruses depend for survival in nature on an assistor virus, which is always a member of the family Luteoviridae (referred to here as a 'luteovirus'). For transmission between plants, CP of luteovirus forms aphidtransmissible hybrid virus particles, encapsidating umbraviral RNA (for review, see Taliansky et al., 2000). In nature,...