Barley yellow dwarf viruses (barley yellow dwarf)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Means of Movement and Dispersal
- Seedborne Aspects
- Vectors and Intermediate Hosts
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Barley yellow dwarf viruses
Preferred Common Name
- barley yellow dwarf
Other Scientific Names
- barley yellow dwarf luteoviruses
- cereal yellow dwarf virus
- Hordeum virus nanescens
- maize leaf fleck virus
- red leaf disease of barley
- rice giallume virus
- wheat cereal yellow dwarf virus
International Common Names
- English: BYDV-RPV-isolate; giallume
- Spanish: virus del enanismo amarillo de la cebada
- French: virus de la jaunisse nanisante de l'orge
Local Common Names
- Germany: Haferroete; Weizen Gelbverzwergungsvirus: Gerste
- CYDVR0 (Cereal yellow dwarf polerovirus-RPV)
Taxonomic TreeTop of page
- Domain: Virus
- Unknown: "Positive sense ssRNA viruses"
- Unknown: "RNA viruses"
- Family: Luteoviridae
- Genus: Barley yellow dwarf viruses
Notes on Taxonomy and NomenclatureTop of page
Barley yellow dwarf disease was first reported by Oswald and Houston (1951, 1953). It is caused by a group of luteoviruses named barley yellow dwarf luteoviruses. They are the type member of the luteovirus genus, one of three genera included in the Luteoviridae family (Mayo and Pringle, 1998). The different viruses which cause barley yellow dwarf were originally characterized by Rochow (1969) and Rochow and Muller (1971) who identified five strains from New York state, USA, based on their transmission phenotypes in an experimental system. The nomenclature used to identify the five strains of BYDV reflected the species of aphid that most efficiently transmitted that strain and were not intended to imply absolute specificity. The strains and their principal vectors are MAV (Sitobion avenae), PAV (Rhopalosiphum padi, S. avenae and others), RMV (R. maidis), RPV (R. padi) and SGV (Schizaphis graminum). This nomenclature system has been adopted by nearly all researchers working on barley yellow dwarf. Zhang et al. (1983) reported the strains GPV, DAV and GPDAV from China. Apparently, the Chinese strains have serological relation to the US isolates (MAV, PAV) but differ slightly in the aphid transmission pattern.
Numerous observations support the division of barley yellow dwarf viruses into two viruses and even two genera (Miller and Rasochová, 1997). Currently, the BYDVs are divided in two subgroups based on serology (Waterhouse et al., 1988), cytopathology (Gill and Chong, 1979) and nucleic acid sequences (Martin and D'Arcy, 1990). Group 1 includes PAV, MAV and SGV and group 2 includes RPV and RMV. Group 2 is more closely related to other luteoviruses such as Beet western yellows virus than to group 1 BYDVs (Martin and D'Arcy, 1990). BYDVs have recently been reclassified by the International Committee on Taxonomy of Viruses (ICTV).
DescriptionTop of page BYDVs are included in the Luteovirus genus. Their virus particles are isometric (measuring about 24-28 nm diameter), sedimenting as a single component at ca 104-118 S, and with a buoyant density in CsCl of ca 1.40g/cm³. The protein shell is composed of one polypeptide species of MW 23.5-24.4 x 10³ and the genome consists of one single molecule of positive sense single-stranded RNA (MWs: PAV, 1,850,000; MAV, 2,000,000; RPV, 1,850,000 - 2,000,000) (Waterhouse et al., 1988).
All of the luteovirus nucleotide sequences contain six open reading frames. The MAV and PAV (Group I) nucleotide sequences contain very small ORFs near the 3'-termini of their RNAs that are not present in the nucleotide sequences of the group II viruses (BYDV-RPV, BWYV, PLRV). The nucleotide sequences of the group II luteoviruses contain much larger ORFs near their 5'-termini than are found in the PAV and MAV sequences (Domier, 1995; Miller and Rosochová, 1997).
DistributionTop of page It is not surprising that BYDVs are of global importance (Lister and Ranieri, 1995) because they have a very wide host range in the Poaceae and can be spread efficiently by several aphid vectors that are prevalent worldwide. They are present in most of the countries where their vectors (Rhopalosiphum padi, Sitobion avenae, Metopolophium dirhodum, Rhopalosiphum maidis, Schizaphis graminum among others) have been reported.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Azerbaijan||Present||Mustafayev et al., 2013|
|China||Restricted distribution||EPPO, 2014|
|-Nei Menggu||Present||EPPO, 2014|
|India||Restricted distribution||CMI, 1969; Tandon et al., 1990; EPPO, 2014|
|-Himachal Pradesh||Restricted distribution||EPPO, 2014|
|Iran||Present||Mahlooji and Makoui, 1990|
|Israel||Present||CMI, 1969; von and Wechmar Gera, 1990; EPPO, 2014|
|Japan||Present||CMI, 1969; Kojima, 1997; EPPO, 2014|
|Jordan||Present||CMI, 1969; Makkouk et al., 1990; EPPO, 2014|
|Lebanon||Present||Makkouk et al., 1990|
|Pakistan||Present||CMI, 1969; Aslam and Ahmad, 1990; EPPO, 2014|
|Syria||Present||Makkouk et al., 1990|
|Thailand||Present||Lister and Ranieri, 1995|
|Turkey||Present||Makkouk et al., 1990|
|Egypt||Present||CMI, 1969; Makkouk et al., 1990; EPPO, 2014|
|Ethiopia||Present||Yusuf et al., 1992|
|Kenya||Present||Introduced||1983||Invasive||Wangai, 1990; IPPC-Secretariat, 2005|
|Libya||Present||Makkouk et al., 1990|
|Morocco||Present||El Yamani & Hill, 1990|
|Mozambique||Present||Lister and Ranieri, 1995|
|South Africa||Present||CMI, 1969; von Wechmar, 1990; EPPO, 2014|
|Tanzania||Present||Lister and Ranieri, 1995|
|Tunisia||Present||Makkouk et al., 1990|
|Zimbabwe||Present||Lister and Ranieri, 1995|
|-Alberta||Present||CMI, 1969; Haber, 1990; EPPO, 2014|
|-British Columbia||Present||CMI, 1969; EPPO, 2014|
|-Manitoba||Present||CMI, 1969; Haber, 1990; EPPO, 2014|
|-New Brunswick||Present||CMI, 1969; EPPO, 2014|
|-Ontario||Present||CMI, 1969; Haber, 1990; EPPO, 2014|
|-Prince Edward Island||Present||CMI, 1969; EPPO, 2014|
|-Quebec||Present||CMI, 1969; Haber, 1990; EPPO, 2014|
|Mexico||Present||CMI, 1969; Ranieri et al., 1993; EPPO, 2014|
|USA||Widespread||CMI, 1969; EPPO, 2014|
|-New York||Present||Gildow, 1990|
|-South Carolina||Present||Gildow, 1990|
|Argentina||Present||de Biasi & de Gurfinkel, 1990|
|Bolivia||Present||Fox et al., 1990|
|Brazil||Present||Caetano, 1984; Ramirez, 1990|
|Colombia||Present||Dubin et al., 1984; Fox et al., 1990|
|Ecuador||Widespread||CMI, 1969; Dubin et al., 1984; Ayala et al., 1997; EPPO, 2014|
|Peru||Present||Dubin et al., 1984; Fox et al., 1990|
|Belgium||Present||CMI, 1969; Signoret and Maroquin, 1990; EPPO, 2014|
|Czech Republic||Restricted distribution||EPPO, 2014|
|Czechoslovakia (former)||Restricted distribution||****||CMI, 1969; EPPO, 2014|
|Denmark||Present||CMI, 1969; EPPO, 2014|
|Finland||Absent, invalid record||CMI, 1969; Peltonen-Sanio & Karjalainen, 1991; EPPO, 2014|
|France||Present||CMI, 1969; Henry et al., 1993; EPPO, 2014|
|Germany||Widespread||****||CMI, 1969; Koch and Huth, 1997; EPPO, 2014|
|Hungary||Restricted distribution||****||CMI, 1969; Pocsai et al., 1995; Papp et al., 1996; EPPO, 2014|
|Ireland||Present||CMI, 1969; Mercer et al., 1989; EPPO, 2014|
|Italy||Widespread||CMI, 1969; Loi et al., 1990; EPPO, 2014|
|Netherlands||Widespread||CMI, 1969; Oswald and Thung, 1955; EPPO, 2014|
|Norway||Widespread||****||CMI, 1969; EPPO, 2014|
|Poland||Present||Hoppe et al., 1983|
|Russian Federation||Restricted distribution||EPPO, 2014|
|-Russia (Europe)||Restricted distribution||EPPO, 2014|
|Spain||Present||Fereres et al., 1989; Comas et al., 1993|
|Sweden||Widespread||****||CMI, 1969; Eweida, 1986; EPPO, 2014|
|Switzerland||Restricted distribution||CMI, 1969; Kobel, 1961; EPPO, 2014|
|UK||Widespread||****||CMI, 1969; Henry et al., 1993; Plumb, 1995; EPPO, 2014|
|Ukraine||Present||Omelchenko & Babayants, 1997|
|Yugoslavia (former)||Present||Tosic et al., 1991|
|Australia||Widespread||CMI, 1969; EPPO, 2014|
|-New South Wales||Present||CMI, 1969; Waterhouse and Helms, 1985; EPPO, 2014; Milgate et al., 2016|
|-Queensland||Present||CMI, 1969; Greber, 1988; EPPO, 2014|
|-South Australia||Present||CMI, 1969; Henry and Francki, 1992; EPPO, 2014|
|-Tasmania||Present||CMI, 1969; Guy et al., 1986; EPPO, 2014|
|-Victoria||Present||CMI, 1969; Sward and Lister, 1987; EPPO, 2014|
|-Western Australia||Present||CMI, 1969; Jones et al., 1990; EPPO, 2014|
|French Southern and Antarctic Territories||Present||Svanella-Dumas et al., 2013|
|New Zealand||Present||CMI, 1969; Johnstone et al., 1990; EPPO, 2014|
Risk of IntroductionTop of page There are no known quarantine regulations for BYDVs because they are not seed transmitted and occur worldwide.
Hosts/Species AffectedTop of page BYDVs are restricted in host range to the Poaceae. Cultivated hosts include all the major cereal crops: barley, maize, oat, rice, rye and wheat (Triticum aestivum, T. durum) (Oswald and Houston, 1951, 1953; Watson and Mulligan, 1960). Many annual and perennial lawn and weed pasture species are also hosts (D'Arcy, 1995). The pasture crops that are mostly affected include ryegrass (Lolium perenne, Lolium multiflorum) (Catherall and Parry, 1987; Eagling et al., 1989), Fescue spp., Bromus spp. (Henry and Dedryver, 1991), cocksfoot (Dactylis glomerata), Phalaris (Phalaris aquatica) and Timothy grass (Phleum pratense) (Guy et al., 1986; Guy, 1988). The known host range of the BYDVs includes more than 150 species (D'Arcy, 1995).
D'Arcy (1995) lists wild Poaceae that have been reported to be naturally infected with BYDVs. These wild hosts may act as reservoirs for the virus. In addition, BYDV has been transmitted under experimental conditions to a wider range of Poaceae (for review, see D'Arcy, 1995).
In general, plants are more sensitive to BYDVs when they are infected at early growth stages. Smith and Sward (1982) showed nearly no damage occurred when the wheat was inoculated when the first node of the stem was visible, compared with up to more than 40% loss when inoculated before tillering. Comeau (1987) suggests that wheat has a temporary rise of resistance at the end of tillering, a period of higher sensitivity during stem elongation and second decrease in sensitivity at flowering time.
The virus spreads systematically throughout the plant.
Growth StagesTop of page Flowering stage, Seedling stage, Vegetative growing stage
SymptomsTop of page Symptoms caused by BYDVs differ with the host species and cultivar, the age and the physiological state of the host plant at the time of infection, the strain and the environmental conditions and can be easily confused with nutritional and abiotic disorders.
Symptoms include leaf discoloration from tip to base and from margin to centre. The discoloration takes on different colours depending on the plant. In barley, the leaf turns bright yellow; in oat, an orange, red or purple discoloration is seen and in wheat, rye and triticale, the infected leaves are generally yellow and sometimes red. In maize, a conspicuous reddening occurs on the lower leaves, while in rice, infected leaves turn yellow to orange (D'Arcy, 1995) Some species of grasses show reddening or yellowing, but many of them are symptomless.
Other leaf symptoms include serrations along leaf borders and corkscrew symptoms as observed on some wheat cultivars infected with RPV-Mexico (M. Henry, unpublished data).
Plants are usually stunted, with a decrease in tiller number and biomass and a weak root system. Suppressed heading, sterility and failure of grains to fill occur in the most severe cases. In the field, symptoms appear usually as yellow or red patches of stunted plants. In hydroponic culture, the root system of BYDV-infected seedlings was initially more severely affected than the shoot, stunting was observed 4 days after infection in roots and only after 18 days in shoots (Hoffman and Kolb, 1997).
In general, PAV causes severe symptoms, MAV moderately severe and RPV, RMV and SGV produce mild symptoms. However, there is a high variability amongst the severity of isolates from the same BYDV strain. Chay et al. (1996) reported PAV isolates ranging from mild to very severe, an RPV isolate producing corkscrew symptoms was isolated in Mexico, and in Ecuador, MAV is known to be extremely severe (Bertschinger, personal communication).
List of Symptoms/SignsTop of page
|Leaves / abnormal colours|
|Leaves / abnormal forms|
|Whole plant / dwarfing|
Biology and EcologyTop of page
BYDVs are not mechanically or seed transmissible, but are transmitted by aphids in a persistent, circulative but non-propagative manner. They are not transmitted to the progeny. Aphids acquire and transmit BYDVs while feeding on the phloem sieve tube elements of host plants. Minimum feeding access times reported for aphids to acquire or inoculate luteoviruses range from 0.1-4.0 h and 0.2-1.0 h, respectively. These reported times include the time required for the aphid's stylets to penetrate to the phloem tissue. Efficient transmission of most luteoviruses requires acquisition (AAP) and inoculation access period (IAP), each of 24 h. The minimum latent period (time from the start of acquisition feeding period until the insect becomes able to infect a plant) is normally between 12 and 24 h (Waterhouse et al., 1988).
After acquisition through the phloem, virions are transported to the aphid hindgut. They cross the hindgut epithelium and are transported in the hemocoel in coated vesicles. The virions must then penetrate the accessory salivary gland basal lamina and plasmalemma to be released into the salivary canals. The virions will then be excreted while the aphid is feeding (Gildow and Gray, 1993). It is suggested that the accessory salivary gland basal lamina possess a selective function that regulates vector specificity through receptors on the plasmalemma and domains of the BYDV virions that interact with these receptors. Van den Heuvel et al. (1994) showed that the aphid protein symbionin was associated with luteovirus transmission and suggested that the interaction between virions and symbionin is involved in maintaining virus integrity and thus specificity.
The specificity of transmission is high. Vector transmission pattern corresponds to serotypes in most cases. However, because more BYDV isolates are being characterized, inconsistencies between transmission pattern and serotypes are becoming more and more common (Power and Gray, 1995). Examples are given by Eweida and Oxelfelt (1985), Lister and Sward (1988), Creamer and Falk (1989) and Halbert et al. (1992).
Many species of aphids infest grasses, including cereal crops. Twenty-five have been reported as vectors of BYDVs (for review, see Halbert and Voegtlin, 1995). The most important vectors include Rhopalosiphum padi, R. maidis, R. rufiabdominalis, Sitobion avenae, Metopolophium dirhodum and Schizaphis graminum.
Interaction between BYDVs
Cross-protection is a phenomenon by which prior-infection by a first (protecting) plant virus prevents or interferes with infection by a second (challenge) virus (Fulton, 1986). Cross-protection exists amongst BYDVs and is correlated with the serological relatedness between the isolates or strains. Wen et al. (1991) showed a high degree of cross-protection between two isolates of BYDV-MAV, moderate cross-protection between BYDV-MAV and BYDV-PAV and no cross-protection between BYDV-PAV, MAV and BYDV-RPV. Generally, there is cross-protection between group I BYDVs but not between group I and II viruses. On the contrary, mixed-infection in a plant with BYDV-PAV and BYDV-RPV can result in a strong symptom aggravation.
A satellite RNA (satRPV RNA) has been associated with an Australian isolate of RPV after glasshouse propagation. This satellite is difficult or impossible to detect in the field and is the only known satellite of a luteovirus. It has no sequence similarity to BYDV genomic RNA and depends on RPV genomic RNA for replication (Miller and Silver, 1991; Silver et al., 1994).
BYDVs are introduced to a new crop by alate aphids originating from infected sources. This is called primary infection. Depending on weather conditions are favourable to aphid development and spread, the virus will spread in the crop (secondary spread). The relative importance of these two phases (primary and secondary infection) is important to consider when deciding on what control strategies to adopt. The primary infection is dependent on the aphid numbers and their ability to transmit the virus. This depends on their species, their previous hosts, the timing of their arrival, the morph (sexual, parthogenetic), their infectivity and their survival and multiplication in the new crops. In different regions of the world and in different climates, reservoirs for the virus have been identified. For example, in western France, maize is an important oversummering reservoir for the aphid and the viruses. The level of infection in this crop and the concordance between autumn flights and sowing of cereals will influence the level of primary infection in the following cereal crops (Henry and Dedryver, 1989). In many regions, pasture grasses and volunteers are important reservoirs. For reviews on epidemiology, see Plumb (1995), Hewings and Eastman (1995), Johnstone (1995) and Irwin and Thresh (1990).
Means of Movement and DispersalTop of page
Twenty-five aphids have been reported as vectors of BYDVs (for review, see Halbert and Voegtlin, 1995). The most important vectors include Rhopalosiphum padi, R. maidis, R. rufiabdominalis, Sitobion avenae, Metopolophium dirhodum and Schizaphis graminum.
Seedborne AspectsTop of page BYDVs are not transmitted through seeds.
Vectors and Intermediate HostsTop of page
|Diuraphis noxia||El-Yamani and Bencharki, 1997.||Insect|
|Metopolophium dirhodum||Rochow, 1970.||Insect|
|Metopolophium festucae||Plumb and Thresh, 1983.||Insect|
|Rhopalosiphum maidis||Rochow, 1970.||Insect|
|Rhopalosiphum padi||Rochow, 1970.||Insect|
|Rhopalosiphum rufiabdominalis||Halbert and Voegtlin, 1995.||Insect|
|Schizaphis graminum||Rochow, 1970.||Insect|
|Sitobion avenae||Halbert and Voegtlin, 1995.||Insect|
|Sitobion fragariae||Vickerman and Wratten, 1979.||Insect|
ImpactTop of page Due to their host range and the widespread occurrence of their vectors, BYDVs are the most economically important virus disease of cereals and the most widespread. Despite this, there are few quantitative estimates of their impact on yield. Recent reviews have been published by Pike (1990) and Lister and Ranieri (1995).
Losses due to barley yellow dwarf can be very serious but vary with the BYDV strains, the growth stage at infection, the wheat varieties and the environmental conditions. In wheat, losses of 47 and 26% have been reported after experimental inoculation with PAV in Kenya (Wangai, 1990) and Mexico (Burnett and Mezzalama, 1992). Losses of around 11-12% due to natural infection have been reported in Morocco (El Yamani and Hill, 1990) and Chile (Ramirez et al., 1992). In Australia, Banks et al. (1995a) reported yield losses of about 2.2 t/ha in a susceptible wheat and about 1.1 t/ha in tolerant varieties.
In barley, yield losses ranging from 3.5 to 29.5% and from 5.6 to 21.1% have been respectively reported in Finland (Rautapaa and Uoti, 1976) and in the USA (Pike, 1990) under natural infection.
In oat, yield losses of 4.4-23.7 and 64.5% were reported from Finland and the USA, respectively (Rautapaa and Uoti, 1976; Martens and Mc Donald, 1970). Recent studies showed a yield reduction due to BYDV-PAV of approximately 4.5% for each 10% increase in barley yellow dwarf incidence (Bauske et al., 1997).
DiagnosisTop of page
BYDVs are distinct from the other viruses infecting cereals in their symptomatology, the morphology and size of their particles and the type of transmission. The symptoms can easily be confused with those of other biotic stresses. Therefore, diagnosis cannot be solely based on symptomatology but must be supported by other techniques such as transmission pattern, serology or PCR.
Before the optimization of serological techniques, transmission tests were the only method for diagnosis of BYDVs. This method is laborious and time-consuming, but sensitive and reliable. It can be used when no other diagnostic methods are available and when it is necessary to define the transmission phenotypes of the isolate. It is the best method to identify an uncharacterized isolate because some of the BYDVs differ from the five main strains described. The detached leaf method has been used by BYDV researchers for diagnosis and BYDV maintenance (Rochow, 1963).
Serologically, BYDV viruses can be divided into subgroups: subgroup I includes PAV, MAV and SGV and subgroup II includes RPV and RMV.
ELISA has been proven to be a fast, sensitive and versatile method to detect plant viruses (Clark and Adams, 1977) including BYDVs (Lister and Rochow, 1979; Rochow and Carmichael, 1979). There is a high serological specificity among BYDVs, so that individual strains can be easily identified. RMV reacts weakly with polyclonal antiserum to RPV, while MAV and SGV cross-react with polyclonal antiserum to PAV. ELISA can be use to detect BYDVs in air-dried samples (Lister et al., 1985) allowing samples to be sent by mail to a testing laboratory.
Monoclonal antibodies have been prepared against several of the BYDVs (Hsu et al., 1984; Diaco et al., 1986; Torrance et al., 1986; Pead and Torrance, 1988; D'Arcy et al., 1990). These can be used to better separate BYDV strains as no cross reaction exists. However, when used in a survey, they might be too restrictive, resulting in some particular isolates with different epitopes not being detected. For example, unusual epitope profiles of BYDVs have been identified in Asia using a range of polyclonal and monoclonal antibodies from various origins (McGrath et al., 1996).
Several private companies sell polyclonal and monoclonal antibodies and ELISA kits for the detection of the five BYDV strains.
Immunosorbent electron microscopy (ISEM) can also be use to detect BYDVs and to separate strains (Forde, 1989), while the tissue-blot immunoassay (TBIA) has been used to evaluate resistance of wheat and barley germplasm to BYDV (Makkouk et al., 1994; Makkouk and Comeau, 1994b).
Detection at the level of a single aphid vector has been achieved by using an amplified ELISA (Torrance, 1987).
Polymerase Chain Reaction (PCR)
Luteovirus-specific PCR primers in RT-PCR (reverse-transcriptase PCR) allow PCR amplification of a 530 bp cDNA fragment from all five strains of BYDV (Robertson et al., 1991). Strains of BYDV can then be distinguished by cutting the PCR products with restriction endonucleases. With the sequence data of more BYDVs being made available, strain specific primers BYDV primers are being developed (French, 1997). BYDV has been detected in single aphids using the RT-PCR technique (Canning et al., 1996).
Rastgour et al. (2005) found that RT-PCR was unable to detect serotypes in crude saps of the infected plants, whereas immunocapture-RT-PCR (IC-RT-PCR) performed more favourably. They concluded that IC-RT-PCR could be routinely and efficiently used for BYDV detection.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.Cultural Practices
Cultural practices that could help reduce BYDVs incidence include changing sowing dates in order to avoid primary infection through viruliferous aphids, removal of cereal regrowths and stubble that can act as reservoirs of virus and vectors and the adoption of adequate cultivation methods (Plumb and Johnstone, 1995). In Australia, delaying sowing might be advisable in winter crops to minimize BYDV-induced grain yield losses. However, the yield benefits need to be balanced against possible yield reductions due to the late sowing (McKirdy and Jones, 1997).
As there is no chemical treatment effective against the virus, chemical control of BYDVs can only be achieved through control of its vectors.
The critical time for control is at an early growth stage (Plumb and Johnstone, 1995). The need for aphid control can either be prophylactic or based on a forecasting system such as those described in Europe by Plumb et al. (1986) and Gillet et al. (1990). Mann et al. (1997) indicated that in England, UK, the spray regimes during spring are of little benefit and that sprays should be applied according to the time of aphid migration relative to crop development and the infectivity of the aphids migrating into the crop. The most commonly used aphicides are organophosphates or synthetic pyrethroids. Imidacloprid, an insecticidal seed treatment, reduced BYD infection under certain conditions (Gray et al., 1996; McKirdy and Jones, 1996). New generation synthetic pyrethroids (alpha-cypermethrin or beta-cyflurin) have been reported to be effective against BYDVs (McKirdy and Jones, 1996).
Success with biological control has been reported from South America, where Sitobion avenae and Metopolophium dirhodum, were controlled through the introduction of Coccinellid predators and Aphelinid and Aphidiid parasites (Zuniga, 1990) and in New Zealand, with the introduction of Aphidius rhopalosiphi (Farrell and Stufkens, 1990). In most areas, natural enemies limit aphid populations and it is important to integrate chemical and natural control methods.
Incorporating resistance or tolerance (Cooper and Jones, 1983) to BYDVs or their vectors is one of the most promising approaches to control. Most of the screening for field 'resistance' to BYD has been directed to the identification of tolerance.
In wheat, sources of tolerance have been reported by several researchers (Burnett et al., 1995). Good tolerance has been reported in germplasm from South America (Ramirez, 1990) and China (M. Henry, unpublished data). Tolerance in the variety Anza (Qualset et al., 1984) has been associated with the presence of the gene Bdv1. It is a partially dominant, partially effective gene that induces slow yellowing (Singh et al., 1993). It is probable that other genes are involved in tolerance to barley yellow dwarf. The winter wheat germplasm lines, Elmo and Caldwell were released as tolerant to BYDV (Ohm et al., 1981; Patterson et al., 1982.
A decrease in virus multiplication has been reported from several wheat relatives, such as Aegilops, Elymus, Elytrigia, Hordeum, Leymus and Thinopyrum (Agropyron) (Sharma et al., 1984, Larkin et al., 1990; Makkouk et al., 1994a; Xu et al., 1994). Recently much effort has been directed toward incorporating these alien-derived resistances into wheat. Resistance is maintained in wheat x Leymus (Plourde et al., 1992), wheat x wheatgrass (Sharma et al., 1989; Goulart et al., 1993), wheat x Agrotricum crosses (Comeau et al., 1994). Thinopyrum intermedium has been widely used to produce resistant introgressed material such as the TC lines (Banks et al., 1995b), Zhong 4 (Xin et al., 1988) and Zhong 5-derived lines (Larkin et al., 1995a). The 42 chromosome winter wheat line P29 and spring wheats TC5, TC6 and TC9 as well as the genetic stock Z1, Z2, Z6 with alien-derived resistance have been registered recently (Banks and Larkin, 1995; Larkin et al., 1995b; Sharma et al., 1997). In TC14, the alien segment is located on 7DL (Hohmann et al., 1996). Tolerance to BYDVs in barley was reported as early as 1961 (Bruehl, 1961; Rochow, 1961). Since then, several lines presenting tolerance to BYDVs have been reported (for review, see Schaller, 1984, Burnett et al., 1995).
The resistance gene Yd2 (Rasmusson and Schaller, 1959) has been used extensively in barley breeding programmes and has been proven to be effective and stable over the years. Cultivars carrying this gene include Atlas 68, CM67 (Schaller and Chim, 1969), Shannon (Vertigan, 1979), Shyri (Vivar et al., 1991), Vixen (Parry and Habgood, 1986), Nomini (Starling et al., 1994) amongst others (Burnett et al., 1995). Delogu et al. (1995) have incorporated the Yd2 gene to high yielding winter wheat. Another resistance gene, yd1, was identified in the cultivar Rojo (Suneson, 1955) but was rarely used in barley breeding programmes because of the low level of resistance it confers.
The gene Yd2 operates by retarding virus multiplication (Jones and Catherall, 1970) and may sometimes loose its effect when placed in a slow growing background. Virus movement from the inoculated leaf towards the roots and subsequently to the growing point was significantly slower in the resistant than susceptible barley genotypes tested by Makkouk et al. (1994b) using ELISA and tissue-printing. Resistance and susceptible lines could be differentiated as early as 3-4 days after inoculation. Yd2 is very effective against the group I BYDVs (PAV, MAV) but only moderately effective against group II (RPV, RMV) (Skaria et al., 1985; Herrera, 1989). In an ICARDA-CIMMYT programme, Yd2 was used extensively with other sources of resistance. Some of the lines show high field tolerance to one or more BYDVs in Ecuador and Mexico (Vivar, personal communication). Chalhoub et al. (1995) identified a Yd2 allele variant that does not originate from Ethiopia and that is overcome by one PAV isolate of BYDV.
Yd2 is located close to the centromere of the long arm of chromosome 3 of barley (Collins et al., 1996). A polypeptide marker of BYDVs resistance identified by Holloway and Heath (1992) can be used as a marker for Yd2. Effort is underway to clone the Yd2 gene.
Sources of tolerance to BYDVs exist in oat but no true resistance has been reported. However, some tolerant lines significantly reduce virus multiplication (Gray et al., 1993) and could qualify as resistant. Many researchers have shown that the tolerance is heritable (Cooper and Sorrels, 1983; Gellner and Sechler, 1986) and that two to four genes contribute to the tolerance (Landry et al., 1984; McKenzie et al., 1985). It appears that released varieties have shown both a good level of protection over a wide range of field conditions and stability over broad geographic areas (Burnett et al., 1995). Selected oat cultivars tolerant to BYD include Otee (Brown and Jedlinsky, 1973); Ogle (Brown and Jedlinsky, 1983) and Hazel (Brown and Kolb, 1989) as the most notable (Burnett et al., 1995). Three major quantitative loci for tolerance to BYDV-PAV have been identified by Jin et al. (1998) to contribute 25, 20 or 17% of the variability.
In addition, good sources of tolerance to BYDVs have been found in other oat species, including Avena sterilis, A. fatua and A. strigosa (Rines et al., 1980; Comeau, 1982, 1984; Jedlinski, 1984).
Good sources of tolerance have been found in triticale (Collin et al., 1990; Burnett and Mezzalama, 1992; Comeau and St-Pierre, 1992). The tolerance has been incorporated into wheat x triticale hybrids (Nkongolo, 1996).
McGrath et al. (1997) transformed oat with the coat protein (CP) genes of BYDV-PAV, BYDV-MAV and BYDV-RPV together with a construct containing the bar gene for herbicide resistance and the uidA reporter gene. Plants with reduced virus titers were found in the T2 (MAV), T3 (RPV) and T4 (PAV) generations. Using the same construct, a few barley plants transformed with CP-PAV showed moderate to high levels of resistance against BYDV-PAV.
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