Resistance to Viruses of Potato and Their Vectors
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Plant Pathol. J. 28(3) : 248-258 (2012) http://dx.doi.org/10.5423/PPJ.RW.06.2012.0075 The Plant Pathology Journal pISSN 1598-2254 eISSN 2093-9280 © The Korean Society of Plant Pathology Mini-Review Open Access Resistance to Viruses of Potato and their Vectors Peter Palukaitis* Department of Horticultural Sciences, Seoul Women’s University, Seoul 139-774, Korea (Received on June 3, 2012; Accepted on July 1, 2012) Potato (Solanum tuberosum) is one of the most important virus-free seed tubers could be produced in areas that were food crops worldwide and yields of potato can be affect- isolated climatically and geographically from both sources ed by virus infection. While more than 40 viruses have of infection and reduced numbers of virus vectors. been found in potato, only nine viruses (potato leafroll However, the various sanitary measures are labor-intensive, virus, potato viruses A, M, S, V, X and Y, potato mop- expensive and difficult to maintain and thus breeding for top virus and tobacco rattle virus) and one viroid (potato resistance may be a preferred option, in the long term. This spindle tuber viroid) have a significant economic impact mini-review describes the major, agronomically-important on potato, worldwide. This review describes the geo- graphical distribution of the most important viruses potato viruses, as well as resistance genes to both these infecting potato and the genes for resistance or tolerance viruses and their vectors, which have been identified and that have been identified against these various infectious deployed. agents. In some cases such resistance genes have been found only in other Solanum species. Few genes for re- Potato Viruses sistance to the vectors of these viruses have been obtain- ed and even fewer have been deployed successfully. About 40 viruses and two viroids can infect potato natural- However, transgenic resistance in potato has been ly. Another 22 viruses and four viroids can infect potato achieved against seven of these disease agents. experimentally. However, of those viruses that infect potato Keywords : potato, potato viruses, resistance, vectors naturally, only nine viruses and one viroid are considered to have a significant economic impact on a worldwide basis (Table 1). The other viruses either are not known to have an Potato (Solanum tuberosum ssp. tuberosum) is the world’s impact or have effects limited to specific locations. Of the fourth major food crop. The modern potato is a self- other 13 viruses, several currently are limited in their compatible outbreeding tetraploid plant derived from distribution to certain areas of Asia, or the Americas and introductions to Europe from South America in 1570. generally are not found in potato, unless alternate hosts for These introductions were of the cultivated ancestral potato their vectors (thrips, leafhoppers and whiteflies) are absent Andigena (S. tuberosum ssp. andigena), another tetraploid, due to drought or other environmental factors. Thus, it is which probably evolved from diploid species. Potato had conceivable that many of these viruses may yet emerge into been adapted to long-day growth conditions by the end of viruses problematic in potato. the 18th century, by selection for early tuber-bearing plants The economic effects of virus infection vary with location, giving high yields. With time, a progressive “degeneration” potato cultivars and other environmental factors. Viruses of the crop was observed, increasing with subsequent such as Potato leafroll virus (PLRV) and Potato virus Y generations. This was recognized in the early 20th century (PVY) can affect yield substantially, with up to 80% losses, as being due to virus infection (Salaman, 1921). Subsequently, while viruses producing mild or latent symptoms, such as it was realized that several viruses were transmitted by Potato virus X (PVX) and Potato virus S (PVS), show yield aphids to potato, and then later still that other vectors trans- losses of at most 10 to 20% (Burton, 1989; Kojima and mitted several viruses to potato. These various observations Lapierre, 1988). led to the introduction of virus disease regulations, the PLRV is probably the most important and damaging development of seed certification schemes and breeding for potato virus on a worldwide basis and is found in all regions virus resistance in a number of countries. Thus, high-grade, where potato is grown. PVY is also widespread and causes considerable damage, depending on the cultivar of potato *Corresponding author. and the strain of virus. PVY is more common than PLRV in Phone) +82-2-970-5614, FAX) +82-2-970-5610 some regions of the world. The other potyviruses of potato, E-mail) [email protected] Potato virus A (PVA) and Potato virus V (PVV), are often Potato Viruses Resistance 249 Table 1. The main viruses of potato Virus species (abbreviation) Genus Transmission Global distribution Potato leaf roll virus (PLRV) Polerovirus Aphid - persistent Worldwide Potato virus Y ( PVY) Potyvirus Aphid - non-persistent Worldwide Potato virus A (PVA) Potyvirus Aphid - non-persistent Worldwide Potato virus V (PVV) Potyvirus Aphid - non-persistent Peru, France, The Netherlands, UK Potato virus X (PVX) Potexvirus Contact Worldwide Potato mop-top virus (PMTV) Pomovirus Spongospora subterranea Andes, North America, China, Japan, Russia, Northern Europe Tobacco rattle virus (TRV) Tobravirus Trichodorid nematodes Europe, China, Japan, New Zealand, North/Central/South America Potato virus S (PVS) Carlavirus Contact (aphid - non-persistent) Worldwide Potato virus M (PVM) Carlavirus Aphid - non-persistent (contact) Worldwide Potato spindle-tuber viroid (PSTVd) Pospiviroid Contact (aphid) Worldwide grouped together with PVY (as “mosaics”) in virus disease virus accumulation, resistance to virus movement, mature regulation schemes, since they are not found as frequently plant resistance, tolerance, and resistance to virus vectors. and usually are not associated with severe disease; PVA can Resistance to infection is a reduction in the number of be found worldwide (except in the Andes) and causes plants that become infected under field conditions. In symptoms varying from mild to severe, while PVV has a resistance to virus accumulation, the plants become infected very limited distribution and until recently was not con- but virus accumulates only to low levels in the plants. In sidered important. PVX occurs worldwide, but generally resistance to virus movement, the virus is impeded in its produces only mild symptoms. Nevertheless, PVX could movement to other parts of the plant, including the tubers. still cause reductions in tuber yield. In particular, PVX is a Mature plant resistance occurs in most cultivars against problem in mixed infections with potyviruses, where a most viruses, but varies with timing, environmental condi- severe synergistic disease is produced. The other four tions and virus strain. Tolerance is a resistance to disease, in viruses, Potato virus M (PVM), PVS, Potato mop-top virus which virus accumulates, but the crop shows little or no (PMTV) and Tobacco rattle virus (TRV) are important, but effect. Most of the specific resistance genes described in only in localized areas, even though some (PVS and PVM) potato specify resistance to viruses in which infection is have a worldwide distribution (Table 1). PMTV is found limited by inducing either an extreme resistance (ER) with primarily in cooler climates, while TRV is found in light little or no virus accumulation at the site of infection and no sandy soils, which favor its nematode vectors. The only movement, or a hypersensitive response (HR), in which a significant viroid in potato, Potato spindle tuber viroid necrotic lesion developed following infection, usually (PSTVd), occurs in various locations, including N. and S. limiting virus infection to the area around the site of America, China and Eastern Europe, but it is not considered inoculation (Solomon-Blackburn and Barker, 2001a, b). indigenous to potato in Western Europe (Table 1). Genes specifying an ER or an HR, listed in Table 2, have All of the above viruses and the viroid are transmitted been identified for resistance against PVY, PVA, PVV, through tubers, while only PSTVd is transmitted through PVX, PVS and PVM. These have been incorporated in true seeds. In addition, PVX, PVS, PVM and PSTVd are potato lines to limited extents (see below). transmitted by contact, TRV is transmitted by Trichodorid nematodes, PMTV is transmitted by the protist Spongospora PLRV subterranea, and PLRV, PVY, PVA, PVV, PVM and some isolates of PVS are transmitted by aphids (Table 1). In some Breeding for resistance to PLRV has been limited by a cases, PSTVd can be transmitted by aphids via co-encapsi- scarcity of sources of complete resistance and by the labor- dation in virions of PLRV (Querci et al., 1997). intensive field screening procedures required to assess the resistance. Sources of resistance to PLRV have been identi- Resistance to potato viruses fied, but few cultivars exhibit strong resistance. Only one resistant cultivar, Pentland Crown, is grown widely in the Barker and Dale (2006) defined six types of resistance to UK. The resistance in Pentland Crown incorporated both various potato viruses. These included resistance to infec- resistance to infection and resistance to virus accumulation. tion (previously known as field resistance), resistance to The partial resistance to virus infection is polygenic (Davidson, 250 Peter Palukaitis Table 2. Host genes for resistance in potato Virus Resistance Type Gene (synonym) Source Reference Potyviruses 1 PVY/PVA/PVV ER Rysto (= R = Ry) S. stoloniferum [1-3] na 2 PVY ER(Y); Rysto (= R ) S. stoloniferum [1,3,4] PVA HR(A); PVV ER(V)/HR(V) rna 3 PVY/PVA ER(Y)/HR(A) Rysto (= R ) S. stoloniferum [1] n1 1 PVY HR Rysto (Ryn = Nysto ) [1,2] n2 5 2 PVY HR Rysto (R = Nysto ) S. stoloniferum [1] 6 PVA ER Nasto (R = Rym) S. stoloniferum [1,2] PVA ER Ra (= Rasto) S. stoloniferum [5] PVA ER/HR Raadg S. tuberosum spp. andigena [6] PVY ER Ryadg S.t.