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.     

Control of subterranean clover red leaf virus in broad bean crops with aphicides

Apterous Rhopalosiphum padi on the first leaf of cereal plants, developed faster and were more fecund at seedling to stem extension of barley, at tillering to earing of oats and at stem extension and earing of wheat. Feeding site on a mature plant also affected the rate of development and fecundity of the aphids; the largest and most fecund developed on the stems of oat and wheat plants. The consequences of these variations in reproductive performance are discussed in relation to the population dynamics of R. padi and its possible pest status in Britain.     

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.     

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.     

Forecasting migration of cereal aphids (Hemiptera: Aphididae) in autumn and spring

The migration of cereal aphids and the time of their arrival on winter cereal crops in autumn and spring are of particular importance for plant disease (e.g. barley yellow dwarf virus infection) and related yield losses. In order to identify days with migration potentials in autumn and spring, suction trap data from 29 and 45 case studies (locations and years), respectively, were set‐off against meteorological parameters, focusing on the early immigration periods in autumn (22 September to 1 November) and spring (1 May to 9 June). The number of cereal aphids caught in a suction trap increased with increasing temperature, global radiation and duration of sunshine and decreased with increasing precipitation, relative humidity and wind speed. According to linear regression analyses, the temperature, global radiation and wind speed were most frequently and significantly associated with migration, suggesting that they have a major impact on flight activity. For subsequent model development, suction trap catches from different case studies were pooled and binarily classified as days with or without migration as defined by a certain number of migrating cereal aphids. Linear discriminant analyses of several predictor variables (assessed during light hours of a given day) were then performed based on the binary response variables. Three models were used to predict days with suction trap catches ≥1, ≥4 or ≥10 migrating cereal aphids in autumn. Due to the predominance of Rhopalosiphum padi individuals (99.3% of total cereal aphid catch), no distinction between species (R. padi and Sitobion avenae) was made in autumn. As the suction trap catches were lower and species dominance changed in spring, three further models were developed for analysis of all cereal aphid species, R. padi only, and Metopolophium dirhodum and S. avenae combined in spring. The empirical, cross‐classification and receiver operating characteristic analyses performed for model validation showed different levels of prediction accuracy. Additional datasets selected at random before model construction and parameterization showed that predictions by the six migration models were 33–81% correct. The models are useful for determining when to start field evaluations. Furthermore, they provide information on the size of the migrating aphid population and, thus, on the importance of immigration for early aphid population development in cereal crops in a given season.     

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 effect of drought stress and temperature on spread of barley yellow dwarf virus (BYDV)

1 The effect of drought stress and temperature on the dispersal of wingless aphids Rhopalosiphum padi (L.) and the pattern of spread of BYDV (barley yellow dwarf virus) within wheat plants in controlled environment chambers was quantified. Combinations of three different drought stress levels, unstressed, moderate and high stress level, and three different temperatures, 5 ± 1 °C, 10 ± 1 °C, and 15 ± 1 °C, were investigated. 2 With increased temperature there was an increase in the mean distance of visited plants from the point of release and in the number of plants visited and infected with BYDV. Drought stress had no effect on mean distance moved by aphids at any temperature or on plants infected with virus at 10 °C and 5 °C. When plants were drought stressed, the numbers of plants visited and infected were greater at 15 °C than at 10 °C and 5 °C. 3 A greater proportion of plants visited by aphids was infected with BYDV when plants were stressed than when not stressed. At 15 °C a greater proportion of these plants was infected than at lower temperatures. There was no difference between treatments in the numbers of aphids present at the end of the experiment. 4 It is concluded that drought stress and temperature are of considerable importance in virus spread.     

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.     

Predation of Rhopalosiphum padi (Homoptera: Aphididae) by polyphagous predatory arthropods during the aphids' pre-peak period in spring barley

Polyphagous predators (e.g. Araneae, Carabidae and Staphylinidae), collected from spring barley fields during 1981-85, were examined by either gut dissection or a R. padi-specific antiserum in enzyme linked immunosorbent assay (ELISA) in order to detect predation of Rhopalosiphum padi during the aphids' establishment and exponential growth phases. Overall 18% of c. 3000 carabids dissected were shown to feed on R. padi during the aphids' pre-peak period. No overall relationship was found between percentage carabids with R. padi in the diet and peak R. padi densities. Relatively high proportions of Bembidion spp. (particularly B. lampros) and Pterostichus cupreus fed on R. padi during the aphids' establishment phase, and proportions of those predator taxa containing R. padi increased with increasing R. padi densities in both high and low aphid density years. P. melanarius and Harpalus rufipes mainly fed on R. padi during the aphids' exponential growth phase. Overall 11% of c. 1350 predators examined in ELISA gave positive reactions to the R. padi antiserum. Relatively high proportions of linyphiid and lycosid spiders were positive throughout the aphid pre-peak period. Several Acari, Opiliones, Trechus spp. (Carabidae), Philonthus spp. (Staphylinidae), Cantharidae and Chilopoda were positive mainly during the aphids' exponential growth phase. Sample sizes were small, however. Very few of the Bembidion spp. tested in ELISA were positive compared with those examined by gut dissection. The maximum period of R. padi protein (antigen) detection in B. lampros was related to temperature, i.e. 8.5 h at 30°C, 20.5 h at 20°C and 34.5 h at 10°C, respectively. It is suggested that the rate of R. padi protein digestion in B. lampros is faster at higher temperatures than the rate of elimination of prey solids from the guts. Several key predators (in this case B. lampros, P. cupreus and linyphiid spiders) which are abundant in spring cereal fields at a time when R. padi emigrants arrive in the crop and which feed on R. padi during the aphids' establishment phase, are identified. The results are compared with those from similar investigations elsewhere with predators of Sitobion avenae and Metopolophium dirhodum.     

Influence of drought on interactions between Rhopalosiphum padi and ground dwelling predators – A mesocosm study

Climate change will lead to extreme droughts, but it is difficult to predict how this will affect crop pests. In particular, it is unclear how interactions between natural enemies and pests will be influenced. In the field, bird cherry-oat aphids (Rhopalosiphum padi (L.)) have been observed to reside close to, or below the ground surface during dry conditions. We hypothesized that this will increase the niche overlap between R. padi and ground-dwelling predators such as carabid beetles and wolf spiders and that aphid numbers will therefore decline during dry conditions. A fully factorial mesocosm experiment was conducted testing the combined effects of drought and predator presence on aphid position and abundance on barley (Hordeum vulgare) plants. In support of our hypothesis, we found that (a) aphids moved below ground during dry conditions, (b) predators reduced aphid numbers, but only during dry conditions, and (c) predators reduced the proportion of aphids below ground in dry conditions. This increased predation effect during dry conditions was, however, compensated for by a corresponding increase in aphid performance on the plants and so the net effect of drought on aphid numbers ended up being neutral. Thus, pests can be affected by drought in complex ways via a combination top-down and bottom-up mechanisms. Predicting how pest populations will be affected by droughts in the future is thus a formidable research challenge.     

Are primary woody hosts ‘island refuges’ for host‐alternating aphids and important for colonization of local cereals?

Only few studies are available dealing with the relation between winter host density and spatial distribution and spring colonization of winter cereals by the host‐alternating cereal aphid species Rhopalosiphum padi and Metopolophium dirhodum. Large‐scale studies in climatically different agroecosystems in Germany from 2004 to 2006 revealed for R. padi and M. dirhodum larger spring/summer populations in landscapes with higher densities of winter hosts. A small‐scale study was performed in winter wheat fields adjacent to a large hedge with several typical winter hosts plants, bird cherry (Prunus padus) and wild rose species (Rosa spp.) to indentify distance effects (0–8, 8–24 and 24–60 m). Weekly measurements of aphid density between May to July showed significantly higher densities of R. padi compared with those of other aphids. Statistical analysis (Tukey–Kramer test and regression analyses) revealed significant gradients from the hedge to the field centre for R. padi and M. dirhodum. In comparative studies, winged R. padi from winter and adjacent summer hosts were genotyped using four microsatellite markers. The results showed that individuals from a certain winter host were not genetically similar with individuals from neighbouring summer hosts; it, therefore, seems that winter host clones did not significantly contribute to population built‐up in cereal fields over short distances. It could be concluded that on a regional scale, the density of sources for early migrants of R. padi is important for colonization intensity of surrounding summer hosts, but that the high local movement intensity and the relative small proportion of aphids that could be analysed in such tracking studies are blurring close spatial relations within short time periods.     

The effect of temperature and wind on the flight activity of cereal aphids

The lower temperature threshold for take-off in Sitobion avenae obtained from an analysis of daily 12·2 m suction trap catches was 16°C. In the laboratory, the take-off threshold for S. avenae was 17·5°C in increasing temperatures, but 19°C when aphids were kept at constant temperatures for a short period of time. The equivalent thresholds were both 20·5°C for Metopolophium dirhodum. Over a period of 16 h no S. avenae took-off from plants at 10°C, 70% at 15°C and all within 16 h at 20°C. It was concluded that suction trap catches can be used to compare the temperature thresholds for take-off of both different species and morphs of a species of aphid. Emigrants of Rhopalosiphum padi, but not of M. dirhodum, took-off at a higher temperature than the alate exules. High winds were found to delay but not inhibit take-off both in the field, and in the laboratory, using both artificial and plant substrates. All aphids eventually flew, even from favourable hosts. It was concluded that adverse weather conditions will delay but not prevent cereal aphid migration in early summer.     

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.     

Influence of endophyte infection, plant age and harvest interval on Rhopalosiphum padi survival and its relation to quantity of N-formyl and N-acetyl loline in tall fescue

Rhopalosiphum padi L. (Homoptera: Aphididae) is sensitive to loline alkaloids present in tall fescue, Festuca arundinacea Shreb., infected with the endophytic fungus, Acremonium coenophialum Morgan-Jones & Gams. Aphid survival was higher on endophyte-free plants regardless of plant age after germination or age of regrowth tissue after clipping. Survival of aphids on endophyte-infected grass was lower on young tissue but increased as plants aged, although it never reached the same level on endophyte-free plants. Both N-formyl and N-acetyl loline increased as uncut or regrowth tissue aged; however, this was influenced by the age of the plant at the initial cut and the clipping frequency. Although even small amounts of loline cause high aphid mortality, the aphids are able to survive on endophyte-infected plants if the tillers have senescing leaves which contain lower amounts of loline. Preference for senescing leaves may help R. padi avoid plant parts containing high amounts of toxic allelochemicals, thus contributing to higher numbers of aphids on older, endophyte-infected plants.     

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.     

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.     

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.     

Effects of elevated atmospheric CO2 concentrations on phloem sap composition of spring crops and aphid performance

Abstract

The concentration and composition of free amino acids and carbohydrates in the phloem sap of wheat and oilseed rape (OSR) and the effects on the performance of aphids (Rhopalosiphum padi and Myzus persicae) were determined under atmospheric carbon dioxide (CO₂) enrichment. The analysis of phloem sap showed that carbohydrates and amino acid levels of the host plants were significantly affected by elevated CO₂ level. Among carbohydrate concentrations in the phloem sap, significant increases were observed in fructose and glucose in spring wheat under CO₂ enrichment, whereas no changes were observed in OSR. These changes in plant chemistry affected the performance of herbivorous insects (i.e. aphids) in varying ways, positively affecting the relative growth rate (RGR) of R. padi in spring wheat and negatively affecting the RGR of M. persicae on OSR.     

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.     

Searching for wheat resistance to aphids and wheat bulb fly in the historical Watkins and Gediflux wheat collections

Insect pests can reduce wheat yield by direct feeding and transmission of plant viruses. Here we report results from laboratory and field phenotyping studies on a wide range of wheat, including landraces from the Watkins collection deriving from before the green revolution, more modern cultivars from the Gediflux collection (north‐western Europe) and modern UK Elite varieties, for resistance to the bird cherry‐oat aphid, Rhopalosiphum padi (Homoptera: Aphididae) and the English grain aphid, Sitobion avenae (Homoptera: Aphididae). A total of 338 lines were screened for R. padi and 340 lines for S. avenae. Field trials were also conducted on 122 Watkins lines to identify wheat bulb fly, Delia coarctata, preference on these landraces. Considerable variation was shown in insect performance among and within different wheat collections, with reduced susceptibility in a number of varieties, but phenotyping did not identify strong resistance to aphids or wheat bulb fly. Field trials showed within collection differences in aphid performance, with fewer aphids populating lines from the Watkins collection. This differs from development data in laboratory bioassays and suggests that there is a pre‐alighting cue deterring aphid settlement and demonstrates differences in aphid preference and performance on older plants in the field compared with seedlings in the laboratory, highlighting the need for phenotyping for aphid resistance at different plant growth stages. No association was identified between performance of the different insect species on individual varieties, potentially suggesting different nutritional requirements or resistance mechanisms.