The barley yellow dwarf virus in Ukraine

Barley yellow dwarf virus (BYDV) has been identified in the Central region of the Ukraine. BYDV infection in winter wheat crops was shown to be highly harmful. There were no winter wheat cultivars being resistent to BYDV and tolerant cultivars occured rarely. In particular seasons BYDV may give rise to widescale epithoties.     

Properties and isolates of barley yellow dwarf virus

Barley yellow dwarf virus is persistently transmitted by a number of aphid species of which three, Rhopalosiphum padi, Sitobion avenae and Metopolophium dirhodum, are common in most years. Other aphids may be locally important. Isolates of the virus differ in their virulence and geographical distribution and are not transmitted equally well by all aphid vectors. Isolates with similar properties are grouped into strains according to their transmission by vectors and their severity. Changes in strain and aphid occurrence from year to year alter the incidence of virus and its effect on yield. These changes emphasize the need for detailed knowledge of cereal aphid biology and epidemiology of BYDV before effective control can be used.     

Evaluation of oat cultivars and lines under infection with barley yellow dwarf virus

Thirteen domestic and foreign oat cultivars and eight breeding lines bred from the University of Illinois were evaluated for resistance to barley yellow dwarf (BYD) using artificial inoculation with Rhopalosiphum padi viruliferous for an isolate of Barley yellow dwarf virus-PAV endemic to Moscow region origin. Cultivar Blaze and six Illinois lines showed the best grain yields under disease pressure that resembled a BYD epidemic.     

Inheritance and effectiveness of genes in barley that condition tolerance to barley yellow dwarf virus

Tolerance to barley yellow dwarf virus (BYDV) in five Ethiopian barley genotypes was conditioned by major genes which apparently occurred at, or near, the same chromosome locus in each genotype. The genes or alleles differed in effectiveness, one providing a high, three an intermediate and one a low level of tolerance. In plants having a slow rate of growth, the effectiveness of the tolerance genes tended to be masked, but the effectiveness of the gene or allele which provided the highest level of tolerance was masked to a lesser extent than that of genes or alleles providing low or intermediate levels of tolerance.     

Concentration of virus and ultrastructural changes in oats at various stages of barley yellow dwarf virus infection

Enzyme-linked immunosorbent assay was used to monitor the concentration of barley yellow dwarf virus (BYDV) in roots and leaves of oats inoculated at the 1 - 2 leaf stage and at the 4 - 5 leaf stage, respectively. Virus was detectable 20 h after inoculation in the roots and after 48 h in the leaves of plants inoculated at the 1 - 2 leaf stage. The virus concentration reached a plateau in the roots after 7–8 days, and was 3–4 times higher than in the leaves. In plants inoculated at the 4 - 5 leaf stage virus was detectable in roots and leaves after 3 and 5 days, respectively. The concentration reached a maximum after 10 days in the roots and after 18 days in the leaves; the concentration in the leaves was 2–3 times higher than in the roots. Virus was readily detectable in seeds from infected plants, both fresh and old dried seeds. However, seed transmission could not be demonstrated. Virus-like particles were first observed in phloem cells of roots 4 days after inoculation, but no ultrastructural changes were detected at this stage. After 5–6 days, disintegrated nuclei and virus-induced vesicles were observed in many cells and abnormal production of callose was found after 10 days. Necrotic phloem cells were observed from day 13, shortly after the appearance of external symptoms.     

The effect of different virus isolates on the expression of tolerance to barley yellow dwarf virus in barley

A barley variety of Ethiopian origin, with a single Mendelian gene con-fering tolerance to barley yellow dwarf virus (BYDV), was equally tolerant to a number of isolates of the virus, whereas a susceptible European barley was more susceptible to isolates transmitted by Rhopalosiphum padi L. than to those transmitted by Macrosiphum (Sitobion) avenae (Fab). However, hybrids between these two varieties homozygous for the Ethiopian tolerance gene were more tolerant to ‘mild’ than to ‘severe’ isolates, irrespective of the vector specificity. The European variety was damaged more severely by all isolates when infected early than when infected late in its development, but the hybrids were damaged more severely by M. awraae-transmitted isolates when infected late. It is suggested that in susceptible plants the concentration, rather than the virulence, of the virus determines disease severity, whereas the reverse is true in plants possessing a gene which reduces virus multiplication. Virus concentration appears to determine the severity of R. padi-transmitted isolates, while virulence determines the severity of M. avenae-transmitted isolates. The latter would also seem to be adapted towards late infection.     

Variation in reaction to barley yellow dwarf virus in ryegrass and its inheritance

Genotypes of Italian and perennial ryegrass differed greatly in their reaction to infection with barley yellow dwarf virus (BYDV). One genotype of perennial ryegrass appeared unaffected, whilst the yield and height of other genotypes were reduced. Progeny of the Italian ryegrass genotypes showed a similar degree of variation. Tolerance, assessed by the effect of the virus on yield, was inherited in a wholly additive manner. Symptoms were rare and their severity was under both additive and non-additive genetic control and strongly correlated with increasing severity of reaction. The utilization of the variation in BYDV reaction in breeding programmes may prove difficult because plants with the highest performance tended to suffer most from BYDV infection, and half-sib progeny ranked in a different order of tolerance in the glasshouse from that of their respective parent genotypes in the field. It is suggested that the observed variation in BYDV response results from a readjustment in the balance of genotypically controlled and environmentally conditioned variation in plant growth.     

The relationship between growth rate and the expression of tolerance to barley yellow dwarf virus in barley

Barley varieties were most tolerant to infection with barley yellow dwarf virus (BYDV) when they grew rapidly, whether the rate of growth was determined by manipulation of the environment or by the innate genetic constitution of the host. A specific, incompletely dominant gene conditioning a high level of tolerance to the virus in certain rapidly growing genotypes in which it occurred naturally, failed to do so when transferred to slower growing genotypes. However, reintroduction into genotypes capable of rapid growth led to full restoration of the gene's effectiveness. Virus-free aphids recovered BYDV less readily from quick- than from slow-growing genotypes, all homozygous for the tolerance gene.     

The effects of barley yellow dwarf virus, aphids and honeydew on Cladosporium infection of winter wheat and barley

Prior infection of both wheat and barley plants by BYDV predisposed their ears to infection by Cladosporium spp. and Verticillium spp. Aphids and honeydew increased the incidence of Cladosporium on wheat ears but not on barley. This difference between crops was attributed to the larger number ot aphids on the wheat. In the glasshouse, aphids and honeydew, but not honeydew alone, increased Cladosporium populations.     

The role of ants in the epidemiology of barley yellow dwarf virus

The effect of ant attendance on the spread of barley yellow dwarf virus by Schizaphis (Toxoptera) graminum (Rondani) was studied under field conditions. Aphids visited by ants multiplied more rapidly than unattended colonies. Secondary spread of virus is caused by apterae which leave overcrowded plants and become established on nearby hosts. Ant-mediated spread of virus is typically confined to the edges of the field. Excavations made by ants around subterranean parts of small grains provide shelter and a suitable feeding site for tht aphid vector.

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Zusammenfassung Die Anregung zur vorliegenden Arbeit wurde dadurch gegeben, daß während einer Schlechtwetterperiode auf unterirdischen Teilen von Hafer- und Gerstenpflanzen von Ameisen besuchte Blattlauskolonien beobachtet wurden. Es handelte sich dabei um die Ameise Lasius neoniger (Emery) und die Blattlaus Schizaphis graminum (Rondani), die als Getreideschädling und als Überträger der Gelbverzwergungsvirose der Gerste in Nordamerika außerordentlich schädlich ist.Die Ergebnisse der Studie bestätigen erneut ältere Beobachtungen, nach denen sich die Populationsdichte der von Ameisen besuchten Blattläuse erheblich erhöht. Verschiedentlich konnte nachgewiesen werden, daß Ameisen Blattlausfeinde, insbesondere Marienkäfer, vertreiben, sobald diese in ihr Territorium eindringen.Die durch Ameisen geförderte Ausbreitung des Virus muß im engen Zusammenhang mit der populationsfördenden Wirkung des Blattlausbesuches gesehen werden. Ungeflügelte Blattläuse verlassen übervölkerte Pflanzen und wandern in die Umgebung der Wirtspflanze ab. Dabei können Entfernungen zurückgelegt werden, die ohne weiteres ausreichen, benachbarte Pflanzen aufzufinden und gegebenenfals mit Virus zu infizieren. Dadurch, daß sich L. neoniger hauptsächlich an den Feldrändern aufhält, bleibt allerdings die Virusausbreitung auf diese Region beschränkt und dringt nicht weiter in den Bestand vor. Selbst unter günstigen Bedingungen, das heißt in der Gegenwart von Ameisen, Blattläusen und Virus während des anfälligen Jugendstadiums des Getreides, bleibt die Ausbreitung des Virus und der dadurch angerichtete Schaden eng lokalisiert.Eine gewisse Bedeutung in der Epidemiologie der Gelbverzwergungsvirose dürfte auch dem Vorkommen von Blattläusen auf unterirdischen Pflanzenteilen zuzuschreiben sein, weil die Läuse hier gegen Witterungs- und Feindeinwirkung geschützt sind.Die von verschiedenen Seiten beobachtete enge Bindung zwischen Ameisen und Blattläusen konnte in dem vorliegenden Fall nicht beobachtet werden. Ein besonderer Instinkt der Ameisen oder Blattläuse scheint mir zur Aufrechterhaltung der Wechselbeziehung nicht notwendig.

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Approved for publication by the Director of the South Dakota Agricultural Experiment Station as Journal Series No. 583.     

Sequence and organization of barley yellow dwarf virus genomic RNA.

The nucleotide sequence of the genomic RNA of barley yellow dwarf virus, PAV serotype was determined, except for the 5'-terminal base, and its genome organization deduced. The 5,677 nucleotide genome contains five large open reading frames (ORFs). The genes for the coat protein (1) and the putative viral RNA-dependent RNA polymerase were identified. The latter shows a striking degree of similarity to that of carnation mottle virus (CarMV). By comparison with corona- and retrovirus RNAs, it is proposed that a translational frameshift is involved in expression of the polymerase. An ORF encoding an Mr 49,797 protein (50K ORF) may be translated by in-frame readthrough of the coat protein stop codon. The coat protein, an overlapping 17K ORF, and a 3'6.7K ORF are likely to be expressed via subgenomic mRNAs.     

Effects of straw disposal and tillage on spread of barley yellow dwarf virus in winter barley

The effects of straw baling or incorporation, combined with ploughing, non-inversion tillage, or direct drilling on the incidence of barley yellow dwarf virus (BYDV), were investigated in plots of winter barley. Virus infection was more prevalent after ploughing (> 40% of the plot area damaged) than after non-inversion tillage (9 – 24%) or direct drilling (< 3%). For each cultivation method, more infection was associated with straw baling than with straw incorporation. There was a strong negative correlation between plot yields and the incidence of BYDV(r= -0.87). Monthly pitfall sampling of non-specific aphid predators showed that the population activity of several taxa (e.g. Linyphiidae, and the larger species of adult and larval Carabidae and Staphylinidae) declined for a short period in autumn-winter following cultivation. The overall treatment-ranking for numbers of predators in autumn (direct drilled > non-inversion > ploughed) was the opposite of subsequent virus infection. The possible roles of aphid predators and other biological mechanisms in determining the observed effects on BYDV, are briefly discussed.     

Vector specificity of barley yellow dwarf virus serotypes and variants in southwestern Idaho

Plants with symptoms of barley yellow dwarf virus (BYDV) obtained in infection feeding assays of aphids collected in the field in Idaho between 1986 and 1988 were tested for virus transmissibility by possible aphid vectors. Isolates obtained during 1987–1988 were also tested with a range of polyclonal antisera which distinguished PAV, MAV, SGV, RPV and RMV serotypes. In 1989 some Idaho (ID) BYDV isolates, maintained as standards for comparison, were serotyped and tested for aphid transmissibility, using 11 species of aphids. There was not always the expected correspondence between serotype and vector specificity for ID isolates. For isolates obtained from field-collected Rhopalosiphum padi, vector transmissibility and serotype corresponded with previous reports; however, 44% of isolates which were serotyped as RMV were also transmissible by species other than Rhopalosiphum maidis. Similarly, the transmissibility of the ID laboratory standards did not always conform to the reported vector specificity of serotypes. The laboratory ID-MAV culture was transmitted by Metopolophium dirhodum and Myzus persicae as well as by Sitobion avenae. The laboratory ID-SGV culture was transmitted by R. padi and 5. avenae as well as by Schizaphis graminum. The ID-RPV culture was transmitted by S. graminum and Rhopalosiphum insertum as well as R. padi. Both of two laboratory ID-RMV cultures were transmissible by R. insertum and R. padi transmitted one of them. The results indicate that, for isolates collected in Idaho, vector specificity cannot be assumed from their serotypes.     

The role of subterranean aphids in the epidemiology of barley yellow dwarf virus

Survival and dispersal of three root aphid species under greenhouse conditions was greatly enhanced in the presence of Lasius neoniger Emery. the ant was observed carrying three root aphid species but not two above-ground species. All attempts to transmit barley yellow dwarf (BYDV) with Forda olivacea Rohwer, Geoica utricularia (Passerini), Anoecia setariae Gillette & Palmer, or Aphis maidiradicis Forbes failed. However, A. maidiradicis was able to transmit cucumber mosaic virus. Rhopalosiphum padi (L.) could acquire BYDV from barley roots, but it rarely infected plants when feeding on roots.

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Zusammenfassung Die vorliegende Arbeit versucht die Bedeutung unterirdisch lebender Blattläuse für die Gelbverzwergungsvirose der Gerste zu klären. Sie schliesst sich damit früheren Untersuchungen über die Beziehung zwischen Schizaphis graminis (Rond.), einem Vektor des Virus, und der Ameise Lasius neoniger Emery an.Es konnte beobachtet werden, dass L. neoniger drei Wurzellausarten wegtrug, die in einer Versuchsarena freigelassen wurden, während zwei oberirdisch-lebende Blattlausarten nicht transportiert wurden. Die Gegenwart von Ameisen im Versuchsareal förderte auch die Vermehrung von Wurzelläusen auf Getreide und deren Ausbreitung auf benachbarte Pflanzen. Diese ausbreitungsfördende Wirkung des Blattlausbesuchs könnte von epidemiologischer Bedeutung sein, falls sich Wurzelläuse als Vektoren erweisen sollten. Alle Versuche, das Virus mit Forda olivacea Rohwer, Geoica utricularia (Passerini), Anoecia setariae Gillette & Palmer oder Aphis maidiradicis Forbes zu übertragen, schlugen jedoch fehl. Das nichtpersistente Gurkenmosaik konnte allerdings durch A. maidiradicis übertragen werden.Neben den typischen Wurzelläusen wurden unterirdisch lebende Rhopalosiphum padi L. im Freiland angetroffen. Übertragungsversuche ergaben, dass das Gelbverzwergungsvirus von der Blattlaus aus den Wurzeln aufgenommen werden konnte, dagegen gelang es nur selten, Getreidepflanzen durch auf Wurzeln saugende Blattläuse zu infizieren. Da aber im Freiland die Mehrzahl der unterirdisch lebenden R. padi auf den leicht zu infizierenden Stengelteilen angetroffen wurden, dürfte dieser relativen Widerstandsfähigkeit der Wurzeln gegenüber der Virusinfektion keine grössere Bedeutung zukommen.

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Approved for publication by the Director of the South Dakota Agricultural Experiment Station as Journal Series No. 672.     

Production of cloned cDNA from a Swedish barley yellow dwarf virus isolate

A cDNA library was produced from the RNA of a Swedish MAV-like isolate of barley yellow dwarf virus (BYDV). The procedure involved random priming and the ds cDNA was cloned into the EcoRl site of the plasmid pUC19. Among the clones obtained some hybridised specifically with MAV-like isolates whereas others also hybridised with PAV-like isolates. Only very weak hybridisation was observed with an RPV-like isolate. An Australian cDNA clone, reported to be PAV-specific (pBY82, Waterhouse, Gerlach & Miller, 1986), hybridised with Swedish MAV-like but not with PAV-like isolates. Probes prepared from the clones detected virus in plant extracts by dot-blot hybridisation with sensitivity greater than that of ELISA. Virus was also readily detected in extracts of viruliferous aphids.     

Transmission of barley yellow dwarf virus by field collected aphids (Homoptera: Aphididae) and their relative importance in barley yellow dwarf epidemiology in southwestern Idaho

Populations of cereal aphids were sampled from 1985–1988 and assayed for transmission of barley yellow dwarf virus (BYDV), Rhopalosiphum padi, Rho-palosiphum maidis, Sitobion avenae, Metopolophium dirhodum, Schizaphis graminum and Macrosiphum euphorbiae collected from host plants transmitted BYDV in bioassays. Of the 1028 Diuraphis noxia collected from plants, one may have transmitted BYDV. The isolate involved resembled SGV in serological and biological characteristics, but since it was not recoverable by any of more than 800 D. noxia subsequently tested, we suspect it may have been a contaminant. Among those aphids collected during the autumn from a suction trap adapted for live collection, R. padi transmitted BYDV most frequently. Other trapped species which transmitted BYDV included: R. maidis, Rhopalosiphum insertum, Macrosiphum euphorbiae, Metopolophium dirhodum and Ceruraphis eriophori. An adapted Infectivity Index indicated that R. padi is by far the most important vector of BYDV during the autumn sowing season in southwestern Idaho. Male R. padi consistently transmitted BYDV more frequently than did females collected during the same period.     

The Yd2 resistance to barley yellow dwarf virus is effective in barley plants but not in their leaf protoplasts

The Yd2 gene for "resistance" to barley yellow dwarf virus (BYDV) has been widely used in barley ( Hordeum vulgare ). We have tested Australian isolates of BYDV of varying severity against barley genotypes with and without the Yd2 gene and report here a positive relationship between symptoms and virus levels determined by ELISA. Cultivar Shannon is the result of backcrossing the resistant line CI 3208 to cultivar Proctor, a susceptible line. It appears to be intermediate in reaction to BYDV between Proctor and CI 3208, although it carries the major gene, Yd2. Unlike the whole plant studies, no significant differences were observed with regard to the ability of protoplasts derived from these various genotypes to support BYDV replication. It is therefore demonstrated for the first time that the Yd2 gene is not among the small number of resistance genes which are effective against virus replication in isolated protoplasts.     

Species composition of aphid vectors (Hemiptera: Aphididae) of barley yellow dwarf virus and cereal yellow dwarf virus in Alabama and western Florida

Yellow dwarf is a major disease problem of wheat, Triticum aestivum L., in Alabama and is estimated to cause yield loss of 21-42 bu/acre. The disease is caused by a complex of viruses comprising several virus species, including Barley yellow dwarf virus-PAV and Cereal yellow dwarf virus-RPV. Several other strains have not yet been classified into a specific species. The viruses are transmitted exclusively by aphids (Hemiptera:Aphididae). Between the 2005 and 2008 winter wheat seasons, aphids were surveyed in the beginning of each planting season in several wheat plots in Alabama and western Florida Collected aphids were identified and bioassayed for their yellow dwarf virus infectivity. This survey program was designed to identify the aphid species that serve as fall vectors of yellow dwarf virus into winter wheat plantings. From 2005 to 2008, bird cherry-oat aphid, Rhopalosiphum padi (L.); rice root aphid, Rhopalosiphum rufiabdominale (Sasaki); and greenbug, Schizaphis graminum (Rondani), were found consistently between October and December. The species of aphids and their timing of appearance in wheat plots were consistent with flight data collected in North Alabama between 1996 and 1999. Both R. padi and R. rufiabdominale were found to carry and transmit Barley yellow dwarf virus-PAV and Cereal yellow dwarf virus-RPV. The number of collected aphids and proportion of viruliferous aphids were low. Although this study has shown that both aphids are involved with introduction of yellow dwarf virus to winter wheat in Alabama and western Florida, no conclusions can be made as to which species may be the most important vector of yellow dwarf virus in the region.