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.     

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.     

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.     

Aerial flow of barley yellow dwarf viruses and of their vectors in western France

During the years 1989–1992 cereal aphids were caught alive in a low level (1.5 m high) suction trap operated in Le Rheu (Brittany, France) and tested for BYDV transmission. In most cases comparisons with data collected simultaneously by a 12.2 m suction trap operating in the same site resulted in good relationships between weekly catches at both heights. Results from transmission tests showed that: (i) the two main BYDV vectors were Rhopalosiphum padi and Metopolophium dirhodum during the years of experiment; (ii) PAV and MAV were the commonest viruses and RPV was relatively scarce; (iii) during spring M. dirhodum appeared to be the most important MAV vector and nearly as good a PAV vector as R. padi; (iv) during autumn R. padi was the only vector of the three viruses with mixed transmission allowing it to transmit also MAV probably by heteroencapsidation. To give an indication of the risk of infection, infectivity indices were calculated by multiplying the numbers of aphids caught by the 12.2 m suction trap by the proportion that were infective. These infectivity indices agreed with field records of primary infections.     

Hydroxamic acids affecting barley yellow dwarf virus transmission by the aphid Rhopalosiphum padi

2,4-Dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA), a hydroxamic acid (Hx) occurring in wheat, was shown to deter feeding by the aphid Rhopalosiphum padi (L.), and to reduce BYDV transmission to the plant. Dual choice tests with wheat leaves showed the preferential settlement of aphids on leaves with lower levels of DIMBOA. Electric monitoring of aphid feeding behaviour showed that in seedlings with higher DIMBOA levels fewer aphids reached the phloem and they needed longer times to contact a phloem vessel than in those with lower levels. When aphids carrying BYDV were allowed to feed on wheat cultivars with different DIMBOA levels, fewer plants were infected with BYDV in the higher DIMBOA cultivars than in the lower ones. Preliminary field experiments showed a tendency for wheat cultivars with higher Hx levels to be more tolerant to infection by BYDV than lower Hx level ones.     

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.     

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.     

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.     

Assessing the risk of primary infection of cereals by barley yellow dwarf virus in autumn in the Rennes basin of France

In the Rennes basin, Rhopalosiphum padi is anholocyclic and represents more than 90% of suction trap catches of potential vectors of barley yellow dwarf virus (BYDV) during autumn. From 1983 to 1987 the possibility of predicting the risk of BYDV infection of batches of barley test seedlings (sampling units) exposed each week from September to December near a 12.2 m high suction trap was investigated. Three kinds of variables were checked as possible predictors: weekly mean or maximum temperatures; weekly catches of R. padi (including or excluding males); and percentage of sampling units infested by aphids. Three contrasting examples were observed: during the first three years (1983–1985), infection was high and its change with time followed temperature, aphid catches and plant infestation changes; in 1986, high numbers of aphids caught and a high proportion of plants infested resulted in only low infection and in 1987, both infestation and infection were very low. Simple linear regression analysis showed that the more reliable predictors of infection were the proportion of infested plants and to a lesser extent the numbers of trapped aphids. Multiple linear regressions including either of the three groups of ‘predicting’ variables did not result in any improvement in the prediction. At a practical level, the use of counts of aphid catches would seem a better compromise between accuracy and consistency of prediction and ease of gathering data than that of plant infestation but any significant improvement of the prediction should be sought in an early estimate of the amount of virus available to aphids before they colonise the plants.     

Winter wheat (Triticum aestivum) response to Barley yellow dwarf virus at various nitrogen application rates in the presence and absence of its aphid vector,Rhopalosiphum padi

Barley yellow dwarf (BYD) is one of the most common diseases of cereal crops, caused by the phloem‐limited, cereal aphid‐borne Barley yellow dwarf virus (BYDV) (Luteoviridae). Delayed planting and controlling aphid vector numbers with insecticides have been the primary approaches to manage BYD. There is limited research on nitrogen (N) application effects on plant growth, N status, and water use in the BYDV pathosystem in the absence of aphid control. Such information will be essential in developing a post‐infection management plan for BYDV‐infected cereals. Through a greenhouse study, we assessed whether manipulation of N supply to BYDV‐infected winter wheat, Triticum aestivum L. (Poaceae), in the presence or absence of the aphid vector Rhopalosiphum padi L. (Hemiptera: Aphididae), could improve N and/or water uptake, and subsequently promote plant growth. Similar responses of shoot biomass and of water and N use efficiencies to various N application rates were observed in both BYDV‐infected and non‐infected plants, suggesting that winter wheat plants with only BYDV infection may be capable of outgrowing infection by the virus. Plants, which simultaneously hosted aphids and BYDV, suffered more severe symptoms and possessed higher virus loads than those infected with BYDV only. Moreover, in plants hosting both BYDV and aphids, aphid pressure was positively associated with N concentration within plant tissue, suggesting that N application and N concentration within foliar tissue may alter BYDV replication indirectly through their influence on aphid reproduction. Even though shoot biomass, tissue N concentration, and water use efficiency increased in response to increased N application, decision‐making on N fertilization to plants hosting both BYDV and aphids should take into consideration the potential of aphid outbreak and/or the possibility of reduced plant resilience to environmental stresses due to decreased root growth.     

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.     

Feeding responses of two grain aphids to barley yellow dwarf virus-infected oats

The probing behavior of greenbugs, Schizaphis graminum (Rondani), and oat-bird cherry aphids, Rhopalosiphum padi (L.), was electronically monitored on healthy oats, Avena sativa (L.), and on oats infected with the RPV-NY isolate of barley yellow dwarf virus (BYDV). S. graminum fed better on infected oats than on healthy oats. This was manifested by a shorter time before initiation of committed phloem ingestion, and by increased duration of ingestion from phloem of infected compared to healthy plants. In addition, on infected oats S. graminum made fewer interruptions in their probing once their stylets were inserted into tissues, including phloem. R. padi fed similarly on infected and healthy oats, except that these aphids made fewer short probes to the phloem (lasting <15 min) on infected compared to healthy oats. BYDV infection of oats increased the rate of population growth of S. graminum relative to that on healthy oats but had no effect on the population growth of R. padi. The proportion of aphids of R. padi which developed into alatae on BYDV-infected oats was significantly greater than on healthy oats, but S. graminum showed no such response.

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Résumé Le comportement de sondage de S. graminum (Rond.) et de R. padi (L.) a été suivi électroniquement sur avoine (Avena sativa L.) saine ou contaminée par l'extrait RPV-NY du virus jaune du nanisme de l'orge (BYDV). S. graminum s'est mieux alimenté sur avoine contaminée que saine; ceci se traduisait par un temps de latence inférieur avant l'ingestion de phloème et par une prolongation de la période d'ingestion. De plus, sur avoine contaminée, S. graminum a moins souvent interrompu le sondage, une fois que les stylets avaient étè insérés dans les tissus, y compris le phloème. Les résultats avec R. padi présentaient les mêmes tendances, mais les améliorations des performances alimentaires de cette espèce sur avoine contaminée ont été moins nettes que pour S. graminum. La contamination de l'avoine par BYDV a accru le taux de croissance de la population de S. graminum par rapport à ce qui a été observé sur avoine saine; l'effet était moins prononcé pour R. padi. La proportion de pucerons devenant aliés était plus élevée sur avoine contaminée que saine, rien de tel n'a été observé avec S. graminum.

     

Characterisation of bird cherry-oat aphid (Rhopalosiphum padi L.) behaviour and aphid host preference in relation to partially resistant and susceptible wheat landraces

The bird cherry-oat aphid (Rhopalosiphum padi L.) is a major pest of wheat (Triticum aestivum L.) and can cause up to 30% yield losses. Heritable plant resistance to aphids is both an economically and ecologically sound method for managing aphids. Here we report how the behaviour and performance of R. padi differs on two resistant, one susceptible wheat landrace and a susceptible elite wheat variety. Feeding behaviour differed among the genotypes, with aphids on resistant lines spending longer in the pathway phase and less time phloem feeding. These behaviours suggest that both inter- and intracellular factors encountered during pathway and phloem feeding phases could be linked to the observed aphid resistance. Locomotion and antennal positioning choice tests also revealed a clear preference for susceptible lines. Although feeding studies revealed differences in the first probe indicating that the resistance factors might also be located in the peripheral layers of the plant tissue, scanning electron microscopy revealed no difference in trichrome length and density on the surface of leaves. Aphids are phloem feeders and limiting the nutrient uptake by the aphids may negatively affect their growth and development as shown here in lower weight and survival of nymphs on resistant genotypes and decreased reproductive potential, with lowest mean numbers of nymphs produced by aphids on W064 (54.8) compared to Solstice (71.9). The results indicate that resistant lines markedly alter the behaviour, reproduction and development potential of R. padi and possess both antixenosis and antibiosis type of resistance.     

Odour stimuli affecting autumn migration of Rhopalosiphum padi (L.) (Hemipter: ornoptera)

R. padi gynoparae settling in the canopy of the winter host, Prunus padus, tended to form aggregations. Differences in the degree of exposure to wind, sun, etc., did not satisfactorily explain the aggregated pattern observed. In laboratory experiments, aphids did not distinguish between leaves taken from nearby aggregations and leaves taken far away from aggregations. The results of laboratory olfactometer tests and field trapping studies indicate that R. padi gynoparae produce an aggregation pheromone. Thus the aggregated settling pattern of gynoparae of R. padi can best be explained as a response to an aggregation pheromone.     

Resistance and susceptibility of some wheat lines to bird cherry-oat aphid <Emphasis Type="Italic">Rhopalosiphum padi</Emphasis> (Hemiptera: Aphididae)

Bird cherry-oat aphid, Rhopalosiphum padi (L.), is one of the most serious pests of cereals, with an almost worldwide distribution. A primary screening test was carried out to gauge the resistance or susceptibility of 40 wheat lines to R. padi, and follow-up experiments were conducted to determine the development and fecundity rates of R. padi on eight lines. The forty wheat lines examined were classified into two major classes and four subclasses: A (e.g., ERWYT 87-1) and B (e.g., ERWYT 87-20, ERWYT 87-11), with the highest average number of aphids 14 days after infestation, subclass C (e.g., ERWYT 88-8, ERWYT 87-6, ERWYT 87-4), with the lowest average number of aphids, and subclass D (e.g., ERWYT 88-12, ERWYT 88-13), with intermediate status. Aphid nymph developmental time and survival both differed among the wheat lines. Differences were also observed in the fecundity of R. padi, the intrinsic rate of increase (r _m), and other parameters among the wheat lines tested. The highest and lowest values of r _m were obtained for ERWYT 87-1 and ERWYT 88-8 (0.36 and 0.26 nymphs/female/day, respectively). Three lines (ERWYT 87-4, ERWYT 87-6, ERWYT 88-8) were relatively resistant to R. padi, which could prove useful in the development of IPM programs for this aphid in wheat fields.     

The price of protection:a defensive endosymbiont impairs nymph growth in the bird cherry-oat aphid,Rhopalosiphum padi

Bacterial endosymbionts have enabled aphids to adapt to a range of stressors,but their effects in many aphid species remain to be established.The bird cherry-oat aphid,Rhopalosiphum padi(Linnaeus),is an important pest of cereals worldwide and has been reported to form symbiotic associations with Serratia symbiotica and Sitobion miscanthi L-type symbiont endobacteria,although the resulting aphid phenotype has not been described.This study presents the first report of R.padi infection with the facultative bacterial endosymbiont Hamiltonella defensa.Individuals of R.padi were sampled from populations in Eastern Scotland,UK,and shown to represent seven R.padi genotypes based on the size of polymorphic microsatellite markers;two of these genotypes harbored H.defensa.In parasitism assays,survival of H.defensa-infected nymphs following attack by the parasitoid wasp Aphidius colemani(Viereck)was 5 fold higher than for uninfected nymphs.Aphid genotype was a major determinant of aphid performance on two Hordeum species,a modern cultivar of barley H.vulgare and a wild relative H.spontaneum,although aphids infected with H.defensa showed 16%lower nymph mass gain on the partially resistant wild relative compared with uninfected individuals.These findings suggest that deploying resistance traits in barley will favor the fittest R.padi genotypes,but symbiontinfected individuals will be favored when parasitoids are abundant,although these aphids will not achieve optimal performance on a poor quality host plant.     

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.     

Suitability of the biomass crop Miscanthus sinensis as a host for the aphids Rhopalosiphum padi (L.) and Rhopalosiphum maidis (F.), and its susceptibility to the plant luteovirus Barley Yellow Dwarf Virus

1 The reproductive performance of two aphid pest species, Rhopalosiphum padi and Rhopalosiphum maidis, was investigated on two seedling growth stages of Miscanthus sinensis, rhizomatous M. sinensis‘Giganteus’ and barley. Rhopalosiphum padi was unable to complete its development on Miscanthus. Rhopalosiphum maidis was most fecund on rhizomatous plants compared with seedling stages. 2 The ability of R. maidis to transmit the RPV isolate of Barley Yellow Dwarf Virus (BYDV) to M. sinensis seedlings was further investigated. Following successful transmission, host plant symptomology and the effect of infection on the yield of Miscanthus were investigated. Total above soil biomass was reduced by around 23% following infection. 3 The inability of R. padi to utilize Miscanthus is reviewed in light of this species’ origin and inability to utilize C₄ host plants. 4 The potential pest status of R. maidis on Miscanthus is discussed together with the impact that Miscanthus cultivation could have on the ecology of this aphid species and BYDV in the U.K.