Serological and biological characterisation of isolates of barley yellow dwarf virus in Sweden in 1983

In 1983, cereal plants showing symptoms of barley yellow dwarf virus (BYDV), collected from 15 localities in Sweden, were tested for BYDV using enzyme-linked immunosorbent assay (ELISA). Antisera against two Swedish isolates of BYDV were used, a mild isolate (27/77) transmitted specifically by Sitobion avenae and a severe one (39/78) transmitted mainly by Rhopalosiphum padi. No virus was detected in 57 of 607 plants of oats and barley tested. Of the 550 plants in which virus was detected, 366 were infected with viruses similar to isolate 27/77, 116 with viruses similar to 39/78 and the remaining 68 reacted strongly with both antisera. When tested, the latter isolates were shown to be mixtures. Thirty-nine selected samples were also tested with antisera against the USA isolates RPV, RMV, MAV and PAV, and for transmission by S. avenae and R. padi. Twenty-six of these samples were transmitted specifically by S. avenae, one was transmitted only by R. padi and the remaining 12 samples were shown to be infected with a mixture of an S. avenae-specific isolate and one transmitted mainly by R. padi. Antisera against PAV and MAV each detected all isolates tested and the results were very similar to those with the antisera to the 39/78 and 27/77 isolates, respectively. None of the field isolates reacted with antisera against RMV or RPV. It was concluded that 1983 was an epidemic year for BYDV in Sweden and that isolates specifically transmitted by S. avenae predominated. Symptoms of infection by these isolates on oat plants ranged from mild to severe.     

Spread of barley yellow dwarf virus and relative importance of local aphid vectors in central Washington

Data from bioassays of field collected aphids, barley indicator plants exposed to natural conditions, and various types of aphid traps were used to describe the spread of barley yellow dwarf virus (BYDV) in wheat and barley near Prosser, Washington. Bioassays were also used to assess the relative importance of local vector species. Of alate aphids collected from grain in the 1982 and 1983 fall migration seasons, 3.4–14–5% transmitted BYDV. Data from concurrent and post-migration assays of resident aphids (apterae and nymphs) reflected an increase in the proportion of infected plants in the field. Maximum increase in the percentage of viruliferous aphids occurred in late November and December of 1982 and November of 1983. The 1982 increase occurred after aphid flights had ceased for the year, suggesting active secondary spread. Collections in pitfall traps and infected trap plants from November to February confirmed aphid activity and virus spread. Rhopalosiphum padi was the most important vector in central Washington in 1982 and 1983 because of its abundance and relative BYDV transmission efficiency. Metopolophium dirhodum was more winter-hardy than R. padi and equal to R. padi in its efficiency as a vector; however, it was not as abundant as R. padi except during the mild winter of 1982–83, when it was a major contributor to secondary spread. Sitobion avenae may be important in years when it is abundant, but it was only a quarter as efficient as R. padi. Rhopalosiphum maidis was a much less efficient vector than R. padi and it only reached high populations in late autumn barley.     

The effects of sowing date and choice of insecticide on cereal aphids and barley yellow dwarf virus epidemiology in northern England

During the mid-1980s, Sitobion avenae became recognised as an important vector of barley yellow dwarf virus (BYDV) in the Vale of York. A field trial at the University of Leeds Farm, North Yorkshire, was carried out during the autumn/winter of 1984-85 to evaluate different control procedures against S. avenae-transmitted BYDV and to investigate its epidemiology. Winter barley was sown on three dates in September, and plots were sprayed with either deltamethrin, demeton-S-methyl or pirimicarb on one of three dates between mid-October and mid-November, making a factorial design. Rhopalosiphum padi, the main vector of BYDV in southern England, were rarely found during the experiment, but the numbers of S. avenae were much higher, reaching a peak of 21% of plants infested in the unsprayed plots of the first sowing date. Single applications of each insecticide reduced populations of S. avenae to zero. Some treatments, particularly in the early sown plots and those treated with pirimicarb, however, did allow some recolonisation, and thus led to increased virus incidence and decreased yields. Sprays applied before the end of the migration of S. avenae were more efficient at controlling BYDV if the insecticide was persistent, otherwise a spray after this period, in November, was more effective. Virus incidence, although reduced by sprays, was generally low in plots sown on 18 and 27 September. In contrast, about 11% of plants were infected in unsprayed plots sown on 6 September and a small yield benefit was obtained with insecticidal treatments. Enzyme-linked immunosorbent assay (ELISA) of plants taken from the plots indicated that MAV- and PAV-like strains were present, and were most likely to have been transmitted by S. avenae.     

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.     

Purification and properties of two Swedish isolates of barley yellow dwarf virus and their serological relationships with American isolates

Two Swedish isolates of barley yellow dwarf virus (BYDV), one (39/78) transmitted much more efficiently by Rhopalosiphum padi than by Sitobion avenae and the other (27/77) transmitted specifically by Sitobion avenae, were purified with yields of 200–300 μg/100 g infected oat leaves. Three light scattering zones were obtained when isolate 39/78 was sedimented in 10–40% (w/w) sucrose density gradients but only one zone with the other isolate. The sedimentation coefficients of 39/78 particles were 94,106 and 150 S, respectively, whereas the 27/77 particles sedimented at 110 S. Four protein bands of mol. wts 31 900, 29 700, 28 000 and 15 000 were detected when disrupted 150 5 particles of the 39/78 isolate were electrophoresed in SDS-polyacrylamide gels. Only one major band of mol. wt 24 500 was detected when 94 or 106 S particles were electrophoresed under the same conditions. The 27/77 isolate yielded one major band of mol. wt 23 500. A weak serological relationship was found between isolates 39/78 and 27/77. In enzyme-linked immunosorbent assay, isolate 39/78 was serologically closely related to New York isolate PAY, whereas 27/77 was closely related to MAY.     

Barley yellow dwarf virus in aphids caught in suction traps, 1969-73

Suction traps operating at low level (1 5 m) were used to catch live alate Rhopalosiphum padi, Macrosiphum (Sitobion) avenae and Metopolophium dirhodum which were tested for transmission of barley yellow dwarf virus (BYDV). The first species caught and infective was R. padi, followed by M. (S.) avenae infective some 2–3 wk later and M. dirhodum 3–4 wk later still. Never more than 11-5% of the annual catch of any species transmitted BYDV and the proportion fluctuated from week to week and between seasons in different years. The relative abundance of infective vectors of ths three species varied; annual numbers of infective M. (S.) avenae and M. dirhodum varied inversely with infective R. padi, the latter also usually transmitted severer virus. The results of the infectivity tests have been compared with the catches of these aphids by the Rothamsted Insect Survey and show that numbers of alate aphids do not necessarily indicate the likely incidence of BYDV.     

Identification, Distribution and Vector Population Dynamics of Barley Yellow Dwarf Virus in Three Cereal-Producing Areas of Spain

ELISA-based surveys during 1985–87 in three major cereal-growing areas of Spain confirmed the presence of barley yellow dwarf virus (BYDV). Samples of small grain cereals and grasses with and without BYDV-like symptoms were collected in the central, southwestern, and northeastern Spain. Infections were found in all cereal species sampled and in some grasses. About 37 % of the samples collected in 1985 were infacted with isolates of the PAV serotype. Isolates of the RPV serotype were less common, and were detected only in samples from the central region at El Encin, Madrid. Only a single sample, collected from El Encin in 1987, was unequivocally diagnosed as containing an isolate of the MAV serotype. Aphid vector population dynamics was monitored during fall and winter of 1984–87 in the central region. Rhopalosiphum padi L. appeared to be the most abundant species during fall and winter months, infesting grasses and volunteer wheat. Other species present were Sitobion avenae (F.), Metopolophium dirhodum (Walker) and Rhopalosiphum maidis (Fitch). Both R. padi and S. avenae seem to be anholocyclic in the central region of Spain, and are able to remain and reproduce on wheat volunteers and grasses until the beginning of spring. S, avenae populations increase quickly on wheat volunteers in April, while populations of R. padi remain low. Therefore, spread of S. avenae-transmitted BYDV types to neighbouring cereal fields seem more likely to occur than spread of other types. Other possible virus reservoirs, such as maize, also need investigation for a better understanding of BYDV epidemiology in the central and other cercal-growing areas of Spain.     

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.     

The effect of increasing dosage of barley yellow dwarf virus on some resistant and susceptible barleys

Five spring barleys, grown either in pots out of doors or in the field, were inoculated with barley yellow dwarf virus (BYDV) using 5, 10, 20 or 50 infective aphids (Rhopalosiphum padi) per plant. Control plants of each variety received no aphids. Infection with all aphid numbers had highly significant adverse effects on all varieties except Cb 1029, an early maturing BYDV-resistant barley of Ethiopian origin. 12583 Co, a locally bred, late maturing barley possessing the same resistance gene as Cb 1029 suffered more in a pot experiment, but less than three susceptible varieties all of which were severely damaged even when few infective aphids were used. Progressive effects with increasing aphid numbers, indicative of dosage response, occurred in some varieties. These effects included delay in heading and increased stunting, but not less yield. In Cb 1029, BYDV infection caused a reduction in the number of heads per plant, but this was partly compensated for by an increase in the number of grains per head. Conversely, BYDV infection in 12583 Co caused an increase in the number of heads, partly offset by a decrease in the number of Brains tier head.     

The effect of barley yellow dwarf virus on field populations of the cereal aphids, Sitobion avenae and Metopolophium dirhodum

The numbers of cereal aphids, especially Metopolophium dirhodum in 1979, and Sitobion avenae in 1980, were significantly increased on BYDV infected wheat and oats in 1979, and wheat, barley and oats in 1980. The differences were probably caused by attraction of alates of each species to virus infected plants which had changed colour as a result of their infection. Significantly more alates of M. dirhodum were found on virus infected oats in 1979, and of S. avenae on oats and barley in 1980, although not on wheat in either year. probably because the colour contrast in wheat was less intense than in the other crops. Flight chamber experiments with alates of both species confirmed their visual attraction to virus-infected leaves. The interaction between virus, vector and host plants is discussed with reference to the ecology of virus spread.     

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.     

Comparison of the potential rate of population increase of brown and green color morphs of Sitobion avenae (Homoptera: Aphididae) on barley infected and uninfected with Barley yellow dwarf virus

Life tables of brown and green color morphs of the English grain aphid, Sitobion avenae (Fabricius) reared on barley under laboratory conditions at 20 ± 1°C, 65% ± 5% relative humidity and a photoperiod of 16 : 8 h (L : D) were compared. The plants were either: (i) infected with the Barley yellow dwarf virus (BYDV); (ii) not infected with virus but previously infested with aphids; or (iii) healthy barley plants, which were not previously infested with aphids. Generally, both color morphs of S. avenae performed significantly better when fed on BYDV‐infected plants than on plants that were virus free but had either not been or had been previously infested with aphids. Furthermore, when fed on BYDV‐infected plants, green S. avenae developed significantly faster and had a significantly shorter reproductive period than the brown color morph. There were no significant differences in this respect between the two color morphs of S. avenae when they were reared on virus‐free plants that either had been or not been previously infested with aphids. These results indicate that barley infected with BYDV is a more favorable host plant than uninfected barley for both the color morphs of S. avenae tested, particularly the green color morph.     

Migration of alate morphs of the bird cherry aphid (Rhopalosiphum padi) and implications for the epidemiology of barley yellow dwarf virus

Suction trapping data indicate three periods of migration of Rhopalosiphum padi in spring, summer and autumn. Four alate morphs are present at different times during the year. A comparison of data from suction traps operating at 12·2 and 1·5 m suggests a different behaviour of females in autumn with more being recorded at 12·2 than 1·5 m. Males, which are only present in autumn, were also more numerous at 12·2 m. During tests to measure barley yellow dwarf virus (BYDV) infectivity, only 9% of female R. padi reproduced on oat seedlings in autumn compared with 74% in summer. Tests on alate female R. padi trapped alive showed that in summer all were exules, but during the first half of September these were largely replaced by gynoparae so that in autumn only 5% of all R. padi trapped at 12·2 m were alate exules. The aerial densities of gynoparae and males were 10 times greater at 12·2 than 1·5 m while densities of alate exules were similar at both heights. It is suggested that gynoparae and males fly higher to increase the chance of finding a taller dispersed host plant. The implications for BYDV epidemiology of the behaviour and presence of the various R. padi alate morphs indicate that autumn-sown cereals emerging before mid-September are particularly at risk from colonisation by alate exules before the transition to a mainly sexual migrant population is complete. Alate exules introduce BYDV from comparatively local sources. The ratio of total R. padi to Sitobion avenae in suction trap samples in autumn usually exceeds 100: 1, but on crops it was only 10: 1. The ratio of alate exule R. padi to S. avenae in suction traps in autumn was only 12: 1, similar to that observed on crops.     

Confirmation of the transmission of barley yellow mosaic virus (BaYMV) by the fungus Polymyxa graminis

Resting spores (cystosori) of Polymyxa graminis, selected from roots of barley plants infected with barley yellow mosaic virus (BaYMV), were used to start mono-fungal sand cultures. Out of 20 attempts using over 800 cystosori, P. graminis became established in 12, and in two of these BaYMV symptoms also occurred. BaYMV was detected by ELISA in extracts of dried roots heavily infected with cystosori and in zoospores of P. graminis. Calculations suggested that, on average, each zoospore carried less than 100 virus particles. In two virus acquisition experiments, non-viruliferous isolates of P. graminis failed to acquire BaYMV from roots of mechanically-inoculated plants. In two further experiments, non-viruliferous isolates were grown on rooted tillers produced from healthy plants and those infected with BaYMV by either vector or mechanical inoculation. Zoospores and cystosori of P. graminis subsequently transmitted the virus, but only from plants where it had been introduced by the vector. Repeated mechanical transmission appeared to have selected a strain of virus that could not be acquired and/or transmitted by the vector. The results provide convincing evidence that P. graminis is a vector of BaYMV but suggest that, in natural populations, only a small proportion of spores may be viruliferous.     

Cereal aphids and the infectivity index for barley yellow dwarf virus (BYDV) in northern England

Suction traps at Leeds University Farm, N. Yorkshire, monitored aerial populations of cereal aphids over three autumns. Different migration patterns were observed between the four main species, Sitobion avenae, Metopolophium dirhodum, Rhopalosiphum padi and R. insertum. The relevance of these patterns to the epidemiology of barley yellow dwarf virus (BYDV) is discussed. Transmission tests revealed S. avenae to be the major vector of BYDV, rather than R. padi, which is responsible for disease outbreaks in the south and west of Britain. An Infectivity Index (II) of 50 has been advocated for R. padi-transmitted BYDV, above which economic damage is likely to occur. This value is shown not to be applicable to the Vale of York, and methods of adapting the data are proposed. Such adjusted II values depend on the behaviour and reproduction of the aphids during the transmission tests, and produce II values that correlate well with levels of field infection in the area.     

Bionomics of aphids reared on cereals and some Gramineae

In controlled temperature, light and relative humidity, Metopolophium dirhodum and Sitobion avenae multiplied more on young Proctor barley than on Blenda oats, and less on Cappelle wheat. Rhopalosiphum padi increased in number fastest on barley and slowest on oats. More survived, and generation lengths seemed shorter, on barley for M. dirhodum and S. avenae and on wheat for R. padi. Tests with young cereals outdoors generally agreed with those in controlled conditions. On mature plants, there were more M. dirhodum on barley, more R. padi on wheat and more S. avenae on oats than on the other cereals. Given a free choice in large cages outdoors, most aphids were found on barley. When allowed to choose between grasses, more M. dirhodum were on Dactylis glomerata, Poa pratensis and Festuca pratensis, more R. padi on Lolium perenne and F. pratensis, and more S. avenae on D. glomerata and L. perenne. Most aphids of all species combined were on F. pratensis, Lolium and Phleum, and fewest on Festuca rubra and Holcus mollis.     

Barley yellow dwarf virus infectivity of alate aphid vectors in west Wales

Live trapping at 0.9 m of alate aphid vectors of barley yellow dwarf virus (BYDV) at Aberystwyth from 1970 to 1979 showed that ten species transmitted the virus to oat test plants. Conversion of percentage infective at 0.9 m to numbers infective based on continuous trapping at 1.2 m showed Rhopalosiphum padi and R. insertum to be the main vector species in most years, whilst Metopolophium dirhodum and Sitobion auenae were normally of minor importance. The data obtained suggest that epiphytotics of BYDV in autumn-sown cereals were caused by numerous infective vectors flying late in the year and transmitting severe strains of the virus. Evidence is presented that gynoparae and males of R. padi are involved in the autumn spread of BYDV and that three further aphid species, Anoecia corni, Metopolophium albidum and M. frisicum are BYDV vectors. The use of live and continuous trapping techniques in forecasting BYDV epiphytotics is discussed.     

The effects of rootstock and scion on tobacco mosaic virus infection in susceptible, tolerant and immune cultivars of tomato

Reciprocal grafts were made between tomato cultivars Potentate, susceptible, and Virocross, tolerant (heterozygous for resistance gene Tm-i) to tobacco mosaic virus (TMV) isolates of Pelham type o and between isogenic lines of cv. Craigella, susceptible and homozygous for gene Tm-i. The grafted plants were inoculated with a type o isolate; both scion and stock inoculation were studied in the former, scion inoculation only, in the latter. With scion inoculation the virus content of a tolerant scion was greater on a susceptible stock than on a tolerant one, but that of a susceptible scion was unaffected by the type of stock: in contrast, the virus content of a tolerant stock was unaffected by the type of scion but that of a susceptible stock was less with a tolerant than with a susceptible scion. With root inoculation the virus contents of both tolerant and susceptible scions were greater on a susceptible than on a tolerant stock. With cv. Craigella the genotype Tm-1/Tm-1 was found to be immune to the type o isolate used, but in grafts the leaves of Tm-1/Tm-1 scions became tolerant to leaf inoculation when on susceptible stocks and the virus entered the stock. Tm-1/Tm-1 stocks became infected when attached to infected, susceptible scions and did not affect the virus content of those scions. The results indicate that a susceptible healthy stock may change the reaction of a tolerant or immune scion to infection by a strain of TMV.     

A novel major gene on chromosome 6H for resistance of barley against the barley yellow dwarf virus

In a mapping population derived from the Ethiopian barley line L94 × Vada, natural infection by barley yellow dwarf virus (BYDV) occurred. While line L94 hardly showed symptoms, Vada was severely affected. The 103 recombinant inbred lines segregated bimodally. The major gene responsible for this resistance mapped to chromosome 6H. We propose to name the locus Ryd3. A subset of recombinant inbred lines, L94, and Vada were planted in a subsequent field test which confirmed the previous field observations. Double antibody sandwich enzyme-linked immunosorbent assays (DAS-ELISA) indicated that the epidemic was due to a combination of the serotypes BYDV-PAV and BYDV-MAV. In the accessions with the least BYDV symptoms no virus was detected, justifying the consideration of the gene as conferring true resistance rather than tolerance to these viruses. In a laboratory/gauze house trial a near-isogenic line carrying the Vada chromosome 6H fragment in an L94 background was affected as much as Vada. The effect of Ryd3 was quantified, and compared with that of the only other known major gene for resistance to BYDV, Ryd2, which is also of Ethiopian origin and is located on chromosome 3H. Both genes seemed to reduce the chance of the viral isolate used in this study to establish infection. In plants in which it became established, the virus concentration reached a similar level as in susceptible accessions, but with less dramatic symptom development. Inoculated plants in which the virus failed to multiply tended to show an increase in the number of ears per plant, resulting in higher grain yield per plant. Ryd3 co-segregates with several PCR-based molecular markers that may serve for marker assisted selection.     

Barley Yellow Dwarf Virus Strain Incidence in Small Grain Cereals in Northeast Spain

Barley yellow dwarf virus (BYDV) was detected in forage cereals and small grain cereals by indirect enzyme-linked immunosorbent assay. Samples of forage cereals collected in the winters of 1987/1988, 1988/1989 and 1989/1990 showed that this crop is a reservoir of BYDV during the end of summer and autumn. PAV-like and MAV-like isolates, in single or mixed infection, were the most common. The proportion of isolates in the infected samples was relatively stable, Samples of winter cereals collected in the springs of 1988, 1989 and 1990 showed that PAV- and MAV-like isolates were widespread. The proportion of samples infected with PAV-like isolates was much more variable than that of MAV-like isolates. The incidence of PAV-like isolates in winter cereals is more dependent on the population of Rhopalosiphum padi during the winter and early spring, than is the incidence of MAV-like isolates on Sitobion avenae density. In northeast Spain (Lleida basin) forage cereals are a constant source of PAV- and MAV- like isolates from which BYDV inoculum is introduced into winter cereals.