The effect of barley yellow dwarf virus on honeydew production by the cereal aphids, Sitobion avenae and Metopolophium dirhodum

Individual S. avenae and M. dirhodum excreted significantly fewer droplets of honeydew on plants infected with BYDV than on healthy plants. S. avenae excreted less honeydew on the ears than on the leaves of wheat. M. dirhodum excreted less than S. avenae on the leaves. The size of honeydew droplets increased with the age of aphids but was not affected by BYDV infection. Possible reasons for the observed effects of BYDV on honeydew excretion are discussed.     

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.     

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.     

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.     

Jasmonate- and salicylate-induced defenses in wheat affect host preference and probing behavior but not performance of the grain aphid,Sitobion avenae

Jasmonate and salicylatemediated signaling pathways play significant roles in induced plant defenses, but there is no sufficient evidence for their roles in monocots against aphids. We exogenously applied methyl jasmonate （MeJA） and salicylic acid （SA） on wheat seedlings and examined biochemical responses in wheat and effects on the grain aphid, Sitobion avenae （Fab.）. Application of MeJA significantly increased levels of wheat＇s polyphenol oxidase, peroxidase and proteinase inhibitor 1, 2 and 6 days after treatment. In twochoice tests, adult aphids preferred control wheat leaves to MeJA or SA treated leaves. Electrical penetration graph （EPG） recordings of aphid probing behavior revealed that on MeJAtreated plants, the duration of aphid＇s first probe was significantly shorter and number of probes was significantly higher than those on control plants. Also total duration of probing on MeJAtreated plants was significantly shorter than on control plants. Total duration of salivation period on SAtreated plants was significantly longer, while mean phloem ingestion period was significantly shorter than on control plants. However, no significant difference in total duration of phloem sap ingestion period was observed among treatments. The EPG data suggest that MeJAdependent resistance factors might be due to feeding deterrents in mesophyll, whereas the SAmediated resistance may be phloembased. We did not observe any significant difference of MeJA and SA application on aphid development, daily fecundity, intrinsic growth rate and population growth. The results indicate that both MeJA and SAinduced defenses in wheat deterred S. avenae colonization processes and feeding behavior, but had no significant effects on its performance.     

Patterns in aphid honeydew production parallel diurnal shifts in phloem sap composition

Variation in phloem sap composition is important in determining aphid performance and is known to occur at both diurnal timescales and in response to plant age. For field grown potato plants, Solanum tuberosum L. (Solanaceae), we determined diurnal variation in components of phloem sap, measured by ethylene diamine tetra‐acetate exudation, and tested for impacts of plant age. The effects of plant age and diurnal cycles on honeydew production by Macrosiphum euphorbiae (Thomas) and Myzus persicae (Sulzer) (both Hemiptera: Aphididae) were also quantified. Both the ratio of sucrose to amino acids and the composition of amino acids in phloem sap varied significantly with time of day. Dietary essential amino acids contributed a smaller proportion of amino acids in the phloem sap of older plants and during early phases of the diurnal cycle. The only significant effect on aphid honeydew production was of the diurnal cycle for Ma. euphorbiae, although increased honeydew production during the day when compared with the production at night, was consistent across the two species. In contrast with studies carried out at seasonal scales, we found limited evidence for variation in phloem sap composition in response to plant age, consistent with our results for honeydew production. These data highlight the need for improved understanding of how seasonal and diurnal physiology of plants influence performance in phloem sap feeding insects.     

Feeding responses of an oligophagous bean aphid, Megoura crassicauda, to primary and secondary substances in Vicia angustifolia

The feeding behaviour of the aphid Megoura crassicauda Mordivilko (Homoptera: Aphididae), which feeds selectively on plants in the genus Vicia (Fabaceae), was studied. The aphids deposited proteinaceous stylet sheaths intercellularly towards the phloem tissues of host plants. Similar stylet sheaths were formed on a Parafilm membrane when host‐specific acylated flavonoid glycosides [two 2″‐O‐(E)‐p‐coumaroyl esters of quercetin 3‐O‐diglycosides] present in the extracts of the narrow vetch, Vicia angustifolia L., were supplied in the solution covered by the membrane. In contrast, their corresponding deacyl analogues, present more abundantly in the host plant tissues, were not stimulatory, which suggested specificity in the structural requirements of the probing stimulants. While the aphids imbibed an artificial diet composed of primary nutrients (e.g., sucrose and amino acids) and produced a large quantity of honeydew, acylated flavonoids alone and non‐acylated flavonoids supplied with the nutrients more or less suppressed honeydew production. These findings implied that the acylated flavonoids serve as a cue to navigate the stylet sheath towards the phloem prior to sap‐sucking, whereas non‐acylated flavonoids may serve as a negative stimulus to refrain from sucking during tissue penetration before tapping the phloem, although the distribution of these compounds in the plant tissues remains unknown. Thus, the feeding behaviour of M. crassicauda appears to be controlled by multiple chemical stimuli in the process of the settling on its host plant.     

Phloem amino acids and the host plant range of the polyphagous aphid, Aphis fabae

This study investigated the relationship between the essential amino acid requirement of the aphid Aphis fabae Scop. and the phloem sap amino acid composition of its host plants. The dietary amino acid requirement of A. fabae varied between clones. One or more of the eight clones of A. fabae tested displayed depressed larval survival, larval growth rate, or r_m on diets lacking histidine, methionine, threonine, and valine, but none of the other five essential amino acids. The required amino acids corresponded closely to the essential amino acids that varied in relative concentrations among 16 plant species tested: histidine, threonine, tryptophan, and valine. It is suggested that the interclonal variation in the dietary requirements of an aphid species may contribute to the intraspecific variation in plant utilisation patterns. The phloem sap amino acid composition and sucrose : amino acid ratio did not differ consistently between host plant species of A. fabae and non‐host species, indicating that phloem amino acid composition is not an important factor in determining the host plant range of this aphid species.     

Relationship between Water Soluble Carbohydrate Content,Aphid Endosymbionts and Clonal Performance of Sitobion avenae on Cocksfoot Cultivars

Aphids feed on plant phloem sap, rich in sugars but poor in essential amino acids. However, sugars cause osmotic regulation problems for aphids, which they overcome by hydrolysing the sugars in their gut and polymerising the hydrolysis products into oligosaccharides, excreted with honeydew. Aphids harbour primary bacterial endosymbionts, which supply them with essential amino acids necessary for survival. They also harbour secondary (facultative) endosymbionts (sfS), some of which have a positive impact on life history traits, although it is not yet known whether they also play a role in providing effective tolerance to differing levels of water soluble carbohydrates (WSCs). We investigated the relationship between WSC content of cocksfoot cultivars and performance of clones of the English grain aphid Sitobion avenae F. We evaluated how clone genotype and their sfS modulate performance on these different cultivars. We therefore examined the performance of genetically defined clones of S. avenae, collected from different host plants, harbouring different sfS. The performance was tested on 10 Dactylis glomerata L. cultivars with varying WSC content. D. glomerata is known as a wild host plant for S. avenae and is also commercially planted. We found that high WSCs levels are responsible for the resistance of D. glomerata cultivars to specific S. avenae clones. The minimum level of WSCs conferring resistance to D. glomerata cultivars was 1.7% dw. Cultivars with a WSC content of 2.2% or higher were resistant to S. avenae and did not allow reproduction. Our results further indicate that sfS modulate to some extend host plant cultivar adaptation in S. avenae. This is the first study revealing the importance of WSCs for aphid performance. Cocksfoot cultivars with a high content of WSCs might be therefore considered for aphid control or used for resistance breeding in this and other grass species, including cereals.     

Performance of clones and morphs of two cereal aphids on wheat plants with high and low nitrogen content

The great variability of the aphid life cycle and their tendency for host alternation gives rise to aphid clones and morphs. Inter‐ and intraclonal variability may be observed in the responses of aphids to various environmental factors. In this study we aimed to evaluate the influence of intrinsic factors (clone and morph type) on the intrinsic rate of increase (r_m) of the English grain aphid, Sitobion avenae (Fabricius), and the bird cherry‐oat aphid, Rhopalosiphum padi (Linnaeus). For each species four apterous clones were collected from established laboratory colonies and compared to assess their relative fitness on high‐ and low‐nitrogen wheat plants under laboratory conditions. The clones had significantly different intrinsic rates of increase on high‐ and low‐nitrogen plants. All R. padi clones had a higher intrinsic rate of increase and mean relative growth rate than S. avenae. Experiments were also conducted to compare the mean fecundity of apterous and alate morphs of S. avenae and R. padi clones on high‐ and low‐nitrogen wheat plants. On high‐nitrogen plants the apterous morphs of S. avenae clones had significantly higher mean fecundity than alate morphs. There were no significant differences between the mean fecundity of alate morphs of the same species on high‐ and low‐nitrogen plants. The results support the idea of better fitness of specific clones/morphs on certain host plants due to higher and lower intrinsic rates of increase.     

Cucurbit aphid‐borne yellows virus (CABYV) modifies the alighting,settling and probing behaviour of its vector Aphis gossypii favouring its own spread

Plant pathogens are able to influence the behaviour and fitness of their vectors in such a way that changes in plant–pathogen–vector interactions can affect their transmission. Such influence can be direct or indirect, depending on whether it is mediated by the presence of the pathogen in the vector's body or by host changes as a consequence of pathogen infection. We report the effect that the persistently aphid‐transmitted Cucurbit aphid‐borne yellows virus (CABYV, Polerovirus) can induce on the alighting, settling and probing behaviour activities of its vector, the cotton aphid Aphis gossypii. Only minor direct changes on aphid feeding behaviour were observed when viruliferous aphids fed on non‐infected plants. However, the feeding behaviour of non‐viruliferous aphids was very different on CABYV‐infected than on non‐infected plants. Non‐viruliferous aphids spent longer time feeding from the phloem in CABYV‐infected plants compared to non‐infected plants, suggesting that CABYV indirectly manipulates aphid feeding behaviour through its shared host plant in order to favour viral acquisition. Viruliferous aphids showed a clear preference for non‐infected over CABYV‐infected plants at short and long time, while such behaviour was not observed for non‐viruliferous aphids. Overall, our results indicate that CABYV induces changes in its host plant that modifies aphid feeding behaviour in a way that virus acquisition from infected plants is enhanced. Once the aphids become viruliferous they prefer to settle on healthy plants, leading to optimise the transmission and spread of this phloem‐limited virus.     

Aphid activities during sieve element punctures

Aphid salivation in sieve elements and phloem sap ingestion were linked to waveforms in the Electrical Penetration Graph (EPG). Non-viruliferousRhopalosiphum padi (L.) (Hemiptera, Aphididae) on barley yellow dwarf virus (BYDV) infected wheat could acquire the virus, which was used as an indication for phloem sap ingestion, whereas virus inoculation by viruliferous aphids on healthy plants was associated with salivation in sieve elements or other phloem cells. Probing was monitored and the waveforms recorded were related to ELISA results of test plants. The EPG patterns A, B, and C are indicative of the stylet pathway phase, whereas patterns E1 and E2 reflect the phloem (sieve element) phase with an unknown activity (E1) or with ingestion and concurrent salivation (E2). Aphids showing pathway and E1 rarely acquired virus, suggesting that little or no phloem sap ingestion can occur during these patterns, whereas those showing additionally pattern E2 did so substantially, indicating phloem sap ingestion. The main pattern related to virus inoculation was E1, although some aphids were able to inoculate plants during pathway. Pattern E1 clearly reflects the most important salivation into sieve elements. Pattern E2 had no clear contribution to virus inoculation, supporting the present hypothesis that during this pattern the saliva is mixed with the phloem sap in the single canal at the stylet tips and ingested immediately, without reaching the plant tissue. Sustained sap ingestion did not affect virus inoculation. So, BYDV inoculation mainly occurs during the first period of a sieve element puncture which is always formed by E1. Implications on persistent virus transmission are discussed.     

Ultrastructural and Immunocytochemical Studies on Effects of Barley Yellow Dwarf Virus – Infection on Fusarium Head Blight, Caused by Fusarium graminearum, in Wheat Plants

The interactions between barley yellow dwarf virus (BYDV) and Fusarium head blight (FHB), caused by Fusarium graminearum, were studied in the two winter wheat cultivars (cvs.), Agent (susceptible to FHB) and Petrus (moderately resistant to FHB), using ultrastructural and immunocytochemical methods. Infections of wheat plants of both cvs. by BYDV increased susceptibility to FHB. BYDV infection caused numerous cytological changes in lemma tissue of both cvs. such as formation of vesicles in the cytoplasm, degradation of fine structures of chloroplasts of both cvs. and accumulation of large starch grains in the chloroplasts. Electron microscopical studies showed that the development of F. graminearum on spike surfaces was not affected in BYDV‐infected plants. After penetration and intercellular growth in lemma tissue, defence responses to Fusarium infections were markedly reduced in BYDV‐diseased plants compared to the tissue of virus‐free plants. At sites of contact of fungal cells with host tissue, depositions of cell wall material were distinctly less pronounced than in tissues of virus‐free plants of cv. Petrus. Detection of β‐1,3‐glucanases and chitinases in lemma tissue of cv. Agent revealed no appreciably increased accumulation of both defence enzymes in F. graminearum‐infected virus‐free and BYDV‐infected tissues compared to the non‐infected control tissue. On the other hand, in cv. Petrus, infection with F. graminearum induced a markedly enhanced activity of both enzymes 3 days after inoculation. The increase of both enzyme activities was less pronounced in BYDV‐infected plants than in tissue exclusively infected with F. graminearum. Cytological studies suggest that in contrast to the susceptible cv. Agent postinfectional defence responses may play still an important role in the resistance of the moderately resistant cv. Petrus to FHB.     

Silencing the expression of the salivary sheath protein causes transgenerational feeding suppression in the aphid Sitobion avenae

Aphids produce gel saliva during feeding which forms a sheath around the stylet as it penetrates through the apoplast. The sheath is required for the sustained ingestion of phloem sap from sieve elements and is thought to form when the structural sheath protein (SHP) is cross‐linked by intermolecular disulphide bridges. We investigated the possibility of controlling aphid infestation by host‐induced gene silencing (HIGS) targeting shp expression in the grain aphid Sitobion avenae. When aphids were fed on transgenic barley expressing shp double‐stranded RNA (shp‐dsRNA), they produced significantly lower levels of shp mRNA compared to aphids feeding on wild‐type plants, suggesting that the transfer of inhibitory RNA from the plant to the insect was successful. shp expression remained low when aphids were transferred from transgenic plants and fed for 1 or 2 weeks, respectively, on wild‐type plants, confirming that silencing had a prolonged impact. Reduced shp expression correlated with a decline in growth, reproduction and survival rates. Remarkably, morphological and physiological aberrations such as winged adults and delayed maturation were maintained over seven aphid generations feeding on wild‐type plants. Targeting shp expression therefore appears to cause strong transgenerational effects on feeding, development and survival in S. avenae, suggesting that the HIGS technology has a realistic potential for the control of aphid pests in agriculture.     

Tripartite Interactions of Barley Yellow Dwarf Virus,Sitobion avenae and Wheat Varieties

The tripartite interactions in a pathosystem involving wheat (Triticum aestivum L.), the Barley yellow dwarf virus (BYDV), and the BYDV vector aphid Sitobion avenae were studied under field conditions to determine the impact of these interactions on aphid populations, virus pathology and grain yield. Wheat varietal resistance to BYDV and aphids varied among the three wheat varieties studied over two consecutive years. The results demonstrated that (1) aphid peak number (APN) in the aphid + BYDV (viruliferous aphid) treatment was greater and occurred earlier than that in the non-viruliferous aphid treatment. The APN and the area under the curve of population dynamics (AUC) on a S. avenae-resistant variety 98-10-30 was significantly lower than on two aphid-susceptible varieties Tam200(13)G and Xiaoyan6. (2) The production of alatae (PA) was greater on the variety 98-10-30 than on the other varieties, and PA was greater in the aphid + BYDV treatment on 98-10-30 than in the non-viruliferous aphid treatment, but this trend was reversed on Tam200(13)G and Xiaoyan6. (3) The BYDV disease incidence (DIC) on the variety 98-10-30 was greater than that on the other two varieties in 2012, and the disease index (DID) on Tam200(13)G was lower than on the other varieties in the aphid + BYDV and BYDV treatments in 2012, but not in 2011 when aphid vector numbers were generally lower. (4) Yield loss in the aphid + BYDV treatment tended to be greater than that in the aphid or BYDV alone treatments across varieties and years. We suggested that aphid population development and BYDV transmission tend to promote each other under field conditions. The aphids + BYDV treatment caused greater yield reductions than non-viruliferous aphids or virus treatment. Wheat varietal resistance in 98-10-30 affects the aphid dispersal, virus transmission and wheat yield loss though inhibits aphid populations from increasing.     

Effect of host‐plant and infection with ‘Candidatus Liberibacter asiaticus’ on honeydew chemical composition of the Asian citrus psyllid,Diaphorina citri

The Asian citrus psyllid (ACP), Diaphorina citri Kuwayama (Hemiptera: Liviidae), transmits the citrus greening pathogen ‘Candidatus Liberibacter asiaticus’ (CLas) by feeding on citrus phloem sap. Because phloem sap is rich in sugars but low in amino acids, ACP sucks large quantities and excretes most of it as honeydew. We studied the chemical composition of ACP honeydew on various host plants. Honeydew samples were analyzed with gas chromatography–mass spectrometry. Fourteen sugars, 13 amino acids, and six organic acids were detected in the honeydew of ACP. Sugars composed about 95% of the total compounds. Sucrose and trehalose were the predominant sugars, composing about 58 and 23% of the total sugars, respectively. Proline, asparagine, aspartic acid, and glutamic acid were the most abundant amino acids in ACP honeydew. The host plant and its infection with CLas had some effect on the honeydew composition. Glucose, chiro‐inositol, myo‐inositol, inositol, maltose, and turanose were lower in honeydew collected from CLas‐infected citrus compared to that collected from non‐infected trees. In CLas‐infected citrus (pineapple sweet orange, Citrus sinensis L. Osbeck) and Bergera koenigii (L.) Spreng. [curry leaf tree (both Rutaceae)] honeydews, valine, alanine, serine, glutamine, glycine, and the organic acids were lower than in honeydew from healthy citrus. Mannose, galactose, inositol, mannitol, an unknown disaccharide, and proline were higher in the honeydew collected from B. koenigii than in honeydew collected from healthy citrus (pineapple sweet orange), whereas fructose, chiro‐inositol, myo‐inositol, trehalose, and lactic acid were lower. The findings of this study help us understand the metabolism and the nutrient needs of ACP that transmits CLas, the pathogen of huanglongbing in citrus.     

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.     

Barley yellow dwarf virus transmission and feeding behaviour of Rhopalosiphum padi on Hordeum bulbosum clones

A Hordeum bulbosum L. (Poaceae) clone A17 was identified, which showed complete resistance to Barley yellow dwarf virus (BYDV) and Cereal yellow dwarf virus (CYDV). It was not possible to infect plants of A17 with BYDV‐PAV, ‐MAV, or with CYDV‐RPV by the aphid vectors Rhopalosiphum padi (L.) or Sitobion avenae (Fabricius) (both Hemiptera: Aphididae). Plants of the A17 clone and of the BYDV‐susceptible H. bulbosum clone A21 revealed some resistance to R. padi compared to the susceptible winter barley cultivar Rubina [Hordeum vulgare L. (Poaceae)]. The development time to the imago was longer and the number of nymphs was reduced on both clones compared with cv. Rubina. The probing and feeding behaviour of R. padi on plants of the H. bulbosum clones was studied over 12 h and compared with that on plants of the barley cv. Rubina. Principal component analysis of the results of the feeding behaviour revealed a clear separation of the H. bulbosum genotypes from Rubina. On H. bulbosum the number of penetrations was higher but total feeding time was shorter. Significant differences were mainly found in the phloem feeding parameters for plants of both clones in comparison to Rubina, with the virus resistant A17 clone having the strongest effect and the susceptible A21 clone being intermediate. Most significant differences were found in parameters of the phloem salivation phase. On A17, an average of less than one (0.9) E1 phase per plant was observed (3.3 on A21 and 5.7 on Rubina) and its duration was reduced to less than 1 min (0.9 min) in comparison to 2.4 min on A21 and 5.7 min on Rubina. Also, the phloem feeding (E2) phase was clearly reduced on A17 plants with 0.5 E2 phases per test and a mean duration of 1.1 min in contrast with 2.9 and 3.5 E2 phases per test and 34.1 and 421.3 min for A21 and Rubina, respectively. These results point towards a phloem‐localized factor for aphid resistance in H. bulbosum, i.e., on A17 plants the phloem salivation time is too short for a successful infection by BYDV leading to vector resistance.     

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.