Feeding behaviour and performance of different populations of the black currant‐lettuce aphid,Nasonovia ribisnigri,on resistant and susceptible lettuce

When crops are bred for resistance to herbivores, these herbivores are under strong selection pressure to overcome this resistance, which may result in the emergence of virulent biotypes. This is a growing problem for crop species attacked by aphids. The Nr‐gene in lettuce confers near‐complete resistance against the black currant‐lettuce aphid, Nasonovia ribisnigri (Mosely) (Hemiptera: Aphididae). Since 2007, populations of N. ribisnigri have been reported in several locations in Europe to infest resistant lettuce varieties that possess the Nr‐gene. The objective of this study was to analyse the behaviour and level of virulence of several N. ribisnigri populations observed to have colonized Nr‐locus‐containing lettuce lines. We analysed the stylet penetration and feeding behaviour, and the performance of these N. ribisnigri populations on resistant and susceptible lettuce lines. Large variation in the degree of virulence to the Nr‐locus‐containing lettuce lines was found among populations of the Nr:1 biotype. The German population was highly virulent on the Nr‐containing resistant lettuce lines, and showed similar feeding behaviour and performance on both the susceptible and resistant lettuces. The French population from Paris was the second most virulent, though reproduction on the resistant lines was reduced. The French population from Perpignan and a population from Belgium, however, showed reduced performance and feeding rate on the resistant compared to the susceptible lettuces. The lettuce background in which the Nr‐gene is expressed influences the level of resistance to the various Nr:1 aphid populations, because the performance and feeding behaviour differed between the aphids on the cultivars (romaine lettuce) compared to the near‐isogenic lines (butterhead/iceberg lettuce). This study also shows that being able to feed on a plant not automatically implies that a population can successfully develop on that plant, because aphids showed phloem ingestion during the 8‐h recording period on resistant lettuce, but were not able to survive and reproduce on the same lettuce line.     

Rearing history affects behaviour and performance of two virulent Nasonovia ribisnigri populations on two lettuce cultivars

Many aphid species have become virulent to host‐plant resistance, which limits the sustainability of insect resistance breeding. However, when this adaptation to resistant plants is associated with fitness costs for the aphids, virulence can be lost in the absence of resistant plants. For two populations of the lettuce aphid, Nasonovia ribisnigri (Mosely) (Hemiptera: Aphididae), we evaluated whether virulence to Nr‐gene‐based resistance was lost on a susceptible lettuce, Lactuca sativa L. (Asteraceae), and assessed possible costs of virulence. The feeding behaviour and performance of these aphids, reared and tested on susceptible and resistant lettuce, were investigated. The rearing plant affected feeding behaviour and performance of the aphids. Temporary reduction and long‐term loss of virulence were found. The total duration of phloem intake was shorter after being reared on susceptible lettuce and tested on resistant lettuce. In addition, one population had a lower survival on resistant lettuce after being reared on susceptible lettuce. There were also indications of fitness costs of the virulence in both populations.     

The effect of co-infestation by conspecific and heterospecific aphids on the feeding behaviour of <Emphasis Type="Italic">Nasonovia ribisnigri</Emphasis> on resistant and susceptible lettuce cultivars

Aphid saliva can suppress the blocking of sieve elements, a reaction that plants employ to inhibit aphid feeding, but aphid saliva can also elicit plant defence responses. Such plant responses might affect interactions between different aphid species and intraspecifically, e.g. among different biotypes. The objectives of our study were to investigate if feeding behaviour and performance of two biotypes of the lettuce aphid Nasonovia ribisnigri are affected by (1) feeding by the other biotype and (2) feeding by the green peach aphid Myzus persicae or the potato aphid Macrosiphum euphorbiae. Additionally the effect of feeding in a group was studied. All experiments were performed on both a resistant and an isogenic susceptible lettuce cultivar. Feeding or probing by conspecific or heterospecific aphids had different effects on Nasonovia ribisnigri biotypes. Aphids were only slightly affected by feeding or probing of the same biotype on both susceptible and resistant lettuce. N. ribisnigri virulent biotype Nr:1 suppressed the resistance against Nr:0 in the resistant cultivar. In contrast, defence was induced by Nr:1 against Nr:0 in susceptible lettuce. Co-infestation by M. euphorbiae and M. persicae had minor effects on Nr:0. Defence against Nr:1 was induced on both susceptible lettuce and resistant lettuce by Nr:0 and M. euphorbiae. Additionally, M. persicae induced defence in resistant lettuce against Nr:1. Effectors in the saliva of Nr:1 aphids are likely responsible for the defence suppression in lettuce. Identification of these effectors could lead to a better understanding of the mechanism of virulence in N. ribisnigri.     

The impact of insecticide resistance in the currant-lettuce aphid, Nasonovia ribisnigri, on pest management in lettuce

Abstract 1 This paper reports on experiments to determine how two different insecticide resistance phenotypes in the aphid Nasonovia ribisnigri (Mosley), which is a major pest of lettuce, change its susceptibility to pyrethroid insecticides and the carbamate pirimicarb. 2 A novel statistical approach determined how the effectiveness of different insecticides was changed by the two resistance phenotypes. This compared the between‐plant distribution of aphid numbers, as opposed to the mean number of aphids per plant. 3 Results from field cage experiments showed that the effect of the resistances differed. Pyrethroid resistance resulted in lower mortality immediately after application of pyrethroids, whereas resistance to pirimicarb shortened the time over which the chemical was effective. 4 The results of laboratory bioassays suggested that these two resistances were not found together in N. ribisnigri. However, the results reported here contradict this assertion. 5 Experiments with insecticide residues showed that reproduction of resistant N. ribisnigri was greater than that of susceptible N. ribisnigri on plants with ageing insecticide residues, even in circumstances where mortality of resistant and susceptible clones of N. ribisnigri were similar. 6 If more than a few aphids are found on a plant then a whole consignment can be rejected for processing. The results reported here suggest that the effect of both insecticide resistances in N. ribsinigri will be to increase the proportion of lettuce heads with an unacceptable number of aphids on them, leading to increased rejection of plants for processing.     

Aphid resistance in Medicago truncatula involves antixenosis and phloem-specific, inducible antibiosis, and maps to a single locus flanked by NBS-LRR resistance gene analogs

Aphids and related insects feed from a single cell type in plants: the phloem sieve element. Genetic resistance to Acyrthosiphon kondoi Shinji (bluegreen aphid or blue alfalfa aphid) has been identified in Medicago truncatula Gaert. (barrel medic) and backcrossed into susceptible cultivars. The status of M. truncatula as a model legume allows an in-depth study of defense against this aphid at physiological, biochemical, and molecular levels. In this study, two closely related resistant and susceptible genotypes were used to characterize the aphid-resistance phenotype. Resistance conditions antixenosis since migratory aphids were deterred from settling on resistant plants within 6 h of release, preferring to settle on susceptible plants. Analysis of feeding behavior revealed the trait affects A. kondoi at the level of the phloem sieve element. Aphid reproduction on excised shoots demonstrated that resistance requires an intact plant. Antibiosis against A. kondoi is enhanced by prior infestation, indicating induction of this phloem-specific defense. Resistance segregates as a single dominant gene, AKR (Acyrthosiphon kondoi resistance), in two mapping populations, which have been used to map the locus to a region flanked by resistance gene analogs predicted to encode the CC-NBS-LRR subfamily of resistance proteins. This work provides the basis for future molecular analysis of defense against phloem parasitism in a plant model system.     

New sources of resistance to lettuce aphids in Lactuca spp.

The use of resistant cultivars is one of the best ways to protect lettuce from aphid pests. At present, there are cultivars available with nearly complete resistance to Nasonovia ribisnigri biotype Nr:0 (based on the Nr gene) and partial resistance to Macrosiphum euphorbiae. Nevertheless, a new biotype of N. ribisnigri (Nr:1) able to overcome the resistance based on the Nr gene is expanding around Europe and has become a major threat of lettuce. In the present work, we report the presence of this new biotype in southeastern Spain, a major lettuce-producing region. Furthermore, a pool of 264 germplasm accessions belonging to Lactuca genus was tested in a greenhouse assay to search for new resistance sources to N. ribisnigri. The most promising accessions were retested in the laboratory to characterize the resistance by means of free-choice and antibiosis assays against biotypes Nr:0 and Nr:1 of N. ribisnigri and against a clone of M. euphorbiae. Three accessions of L. virosa showed resistance against the target aphid species and could be of interest for ongoing breeding programs. The accessions CGN16272 and CGN13361 were both partially resistant to the Nr:1 biotype of N. ribisnigri and to M. euphorbiae, and CGN13355, in spite of not being resistant to N. ribisnigri, showed a near complete resistance to M. euphorbiae. The study of the feeding behavior of N. ribisnigri biotypes showed that the Nr:1 biotype is able to maintain a similar phloem feeding ingestion pattern on genotypes bearing the Nr gene and on N. ribisnigri-susceptible lettuce genotypes. Moreover, as aphids rejected L. virosa as a feeding source due to superficial factors (high level of antixenosis), no differences in the level of antibiosis between such genotypes were detected. A second set of screening assays were conducted on 40 accessions of L. virosa in order to select for resistance against the Nr:1 biotype. The results showed three accessions with high levels of resistance (CGN05148, CGN21399 and CGN16274) against Nr:1 that could be of interest in lettuce breeding programs.     

Foraging ofRhagoletis pomonella flies in relation to interactive food and fruit resources

Feeding behaviour of the lettuce root aphidPemphigus bursarius was monitored electronically on six lettuce varieties. Aphids, monitored for a minimum of 5 h on lettuce roots, performed recognizable feeding behaviour on the susceptible varieties, Webbs Wonderful and Borough Wonder including non probing, pattern A start of new penetration, C (stylet pathway activities) sometimes including clear B-waves (salivary sheath secretions), potential drops (intracellular penetrations), intracellular E (phloem ingestion) distinguishing E₁ and E₂ (Tjallingii, 1990) and occasionally xylem ingestion. Although aphid probing was recorded on the resistant varieties Lakeland and Grand Rapids continued penetration to the phloem elements was deterred and the time spent ingesting phloem was short. Aphids rarely probed the resistant varieties Avoncrisp and Avondefiance and all attempted penetrations quickly terminated.     

Quantifying aphid behavioral responses to environmental change

Aphids are the most common vector of plant viruses, and their feeding behavior is an important determinant of virus transmission. Positive effects of global change on aphid performance have been documented, but effects on aphid behavior are not known. We assessed the plant‐mediated behavioral responses of a generalist aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), to increased CO₂ and nitrogen when feeding on each of three host species: Amaranthus viridis L. (Amaranthaceae), Polygonum persicaria L. (= Persicaria maculosa Gray) (Polygonaceae), and Solanum dulcamara L. (Solanaceae). Via a family of constrained Markov models, we tested the degree to which aphid movements demonstrate preference among host species or plants grown under varying environmental conditions. Entropy rates of the estimated Markov chains were used to further quantify aphid behavior. Our statistical methods provide a general tool for assessing choice and quantitatively comparing animal behavior under different conditions. Aphids displayed strong preferences for the same host species under all growth conditions, indicating that CO₂‐ and N‐induced changes in plant chemistry have minimal effects on host preference. However, entropy rates increased in the presence of non‐preferred hosts, even when preferred hosts were available. We conclude that the presence of a non‐preferred host species affected aphid‐feeding behavior more than changes in plant leaf chemistry when plants were grown under elevated CO₂ and increased N availability.     

Responses of Schizaphis graminum (Homoptera: Aphididae) to leaf excision in resistant and susceptible sorghum

Greenbugs, Schizaphis graminum (Rondani), were reared on intact and excised leaves of varieties of sorghum which differed in their suitability as hosts for this aphid. Aphids grew poorly on intact leaves of three resistant varieties, but grew well on excised leaves of the same varieties. Leaf excision did not affect aphid growth on three susceptible varieties. By electronically monitoring the feeding behaviour of aphids on two resistant and one susceptible variety, significant differences were found in many parameters between aphids assayed on excised vs. intact leaves of only the resistant varieties. Aphids on excised leaves of the resistant varieties, and on excised or intact leaves of the susceptible variety, made fewer probes to the phloem, spending more time ingesting from phloem during each probe, compared to aphids on intact resistant plants. There was a higher level of free amino acids in excised leaves of all varieties, but aphid growth and feeding behaviour improved as a result of excision only on resistant varieties. This observation, coupled with the fact that intact plants of all varieties have similar amino acid levels, indicates that these nutrients are not of primary importance in sorghum suitability to the greenbug. Other explanations for the aphids' responses to excised leaves are discussed.     

The resistance of lettuce (Lactuca sativa L.) toNasonovia ribisnigri: Bionomics ofN. ribisnigri on near isogenic lettuce lines

The intrinsic rate of increase (r_m-values), mean relative growth rates and mortality ofNasonovia ribisnigri (Mosley) (Homoptera, Aphididae) on different lines of lettuce (Lactuca sativa L.) were determined. Near isogenic resistant and susceptible lines, plus their ancestors, were used. Bionomics ofN. ribisnigri on the resistant lines (NrNr) with the dominantNasonovia resistance gene (Nr-gene) differed clearly from the susceptible lines (nrnr). Mortality was high, no larvae reached adulthood, and no reproduction nor honeydew production was seen on the resistant lines. Transfer of aphids to susceptible plants after a period of 2 days on the resistant lines showed no signs of intoxication of aphids. Apparently there is no feeding on the resistant lines. It is not clear whether the aphids can not reach the phloem or do not accept it on the resistant line.     

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.     

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.     

Tissue localisation of lettuce resistance to the aphidNasonovia ribisnigri using electrical penetration graphs

Electrical penetration graphs (EPG's) ofNasonovia ribisnigri (Mosley) (Homoptera, Aphididae) on resistant and susceptible lettuce (Lactuca sativa, Compositae) showed a large reduction in the duration of the food uptake pattern (E2) on the resistant line. No differences in EPG's were observed before the phloem was reached. Therefore, resistance is believed to be located in the phloem vessel. Both mechanical blocking of the sieve element after puncturing and a difference in composition of the phloem sap are possible resistance factors. However, a chemical factor seems more likely because of the specificity of the resistance againstN. ribisnigri.     

Feeding behavior of aphids on narrow‐leafed lupin (Lupinus angustifolius) genotypes varying in the content of quinolizidine alkaloids

Since the beginning of breeding narrow‐leafed lupins [Lupinus angustifolius L. (Fabaceae)] with a low alkaloid content, susceptibility to several aphid species has increased. Therefore, the probing and feeding behavior of Aphis fabae Scopoli, Aphis craccivora Koch, Acyrthosiphon pisum (Harris), Myzus persicae (Sulzer), and the well‐adapted Macrosiphum albifrons Essig (all Hemiptera: Aphididae) was studied over 12 h on narrow‐leafed lupin genotypes containing varying amounts and compositions of alkaloids. We used the electrical penetration graph (EPG) technique to obtain information on the influence of alkaloid content and composition on the susceptibility to various aphid species. Results indicated that the total time of probing of A. fabae, A. craccivora, A. pisum, and M. persicae increased with a reduced alkaloid content, whereas the alkaloid content had no influence on M. albifrons. Almost all of the individuals (>93%) conducted sieve element phases on the highly susceptible genotype Bo083521AR (low alkaloid content). A reduced occurrence of phloem phases was observed during the 12‐h recording on the alkaloid‐rich cultivar Azuro, especially for A. pisum (37.5%) and A. fabae (55.0%). Furthermore, aphids feeding on genotypes with low alkaloid content had in most cases significantly longer sieve element phases than when feeding on resistant genotypes (Kalya: low alkaloid content, yet resistant; Azuro: high alkaloid content, resistant), whereas M. albifrons showed the longest phloem phase on the alkaloid‐rich cultivar Azuro. As most significant differences were found in phloem‐related parameters, it is likely that the most important plant factors influencing aphid probing and feeding behavior are localized in the sieve elements. The aphids’ feeding behavior on the cultivar Kalya, with a low alkaloid content but reduced susceptibility, indicates that not only the total alkaloid content influences the feeding behavior but additional plant factors have an impact.     

Plant stomatal closure improves aphid feeding under elevated CO2

Stomata help plants regulate CO₂ absorption and water vapor release in response to various environmental changes, and plants decrease their stomatal apertures and enhance their water status under elevated CO₂. Although the bottom‐up effect of elevated CO₂ on insect performance has been extensively studied, few reports have considered how insect fitness is altered by elevated CO₂‐induced changes in host plant water status. We tested the hypothesis that aphids induce stomatal closure and increase host water potential, which facilitates their passive feeding, and that this induction can be enhanced by elevated CO₂. Our results showed that aphid infestation triggered the abscisic acid (ABA) signaling pathway to decrease the stomatal apertures of Medicago truncatula, which consequently decreased leaf transpiration and helped maintain leaf water potential. These effects increased xylem‐feeding time and decreased hemolymph osmolarity, which thereby enhanced phloem‐feeding time and increased aphid abundance. Furthermore, elevated CO₂ up‐regulated an ABA‐independent enzyme, carbonic anhydrase, which led to further decrease in stomatal aperture for aphid‐infested plants. Thus, the effects of elevated CO₂ and aphid infestation on stomatal closure synergistically improved the water status of the host plant. The results indicate that aphid infestation enhances aphid feeding under ambient CO₂ and that this enhancement is increased under elevated CO₂.     

Plant water stress intensity mediates aphid host choice and feeding behaviour

1. The effects of drought-induced changes in plant quality on aphid performance and population growth is well-studied. The response of aphid behaviour to plant water limitation has received less attention. Water limitation may affect host-plant colonization by altering the attractiveness of plants. Additionally, plant water limitation may inhibit feeding site establishment and phloem ingestion.
2. Our goal was to examine bird cherry-oat aphid (Rhopalosiphum padi L.) host selection and feeding behaviour under water limitation. We assessed aphid response to well-watered, mildly-stressed, and highly-stressed wheat (Triticum aestivum L.) by evaluating (i) host-plant selection through two-choice assays, (ii) feeding behaviour using the electrical penetration graph technique, and (iii) phloem ingestion by quantifying honeydew production.
3. Aphids were less likely to select highly stressed plants than a mildly stressed or well-watered alternative. Aphids did not distinguish between mildly stressed and well-watered plants. Aphid feeding behaviours, including duration of phloem ingestion, were not affected by water availability. However, honeydew production was reduced under both levels of water limitation. These results suggest that the volume of phloem ingested by aphids per unit time declined on stressed plants. The combination of lower colonization and diminished access to food on stressed plants may lead to a reduction in aphid abundance, independent of the direct effects of nutrition on individual aphid performance.
4. This study highlights the potential contribution of herbivore behaviour to documented changes in aphid abundance on stressed plants and underscores the important role of plant water stress intensity in mediating plant-herbivore interactions.

     

Triticum monococcum lines with distinct metabolic phenotypes and phloem‐based partial resistance to the bird cherry–oat aphid Rhopalosiphum padi

Crop protection is an integral part of establishing food security, by protecting the yield potential of crops. Cereal aphids cause yield losses by direct damage and transmission of viruses. Some wild relatives of wheat show resistance to aphids but the mechanisms remain unresolved. In order to elucidate the location of the partial resistance to the bird cherry–oat aphid, Rhopalosiphum padi, in diploid wheat lines of Triticum monococcum, we conducted aphid performance studies using developmental bioassays and electrical penetration graphs, as well as metabolic profiling of partially resistant and susceptible lines. This demonstrated that the partial resistance is related to a delayed effect on the reproduction and development of R. padi. The observed partial resistance is phloem based and is shown by an increase in number of probes before the first phloem ingestion, a higher number and duration of salivation events without subsequent phloem feeding and a shorter time spent phloem feeding on plants with reduced susceptibility. Clear metabolic phenotypes separate partially resistant and susceptible lines, with the former having lower levels of the majority of primary metabolites, including total carbohydrates. A number of compounds were identified as being at different levels in the susceptible and partially resistant lines, with asparagine, octopamine and glycine betaine elevated in less susceptible lines without aphid infestation. In addition, two of those, asparagine and octopamine, as well as threonine, glutamine, succinate, trehalose, glycerol, guanosine and choline increased in response to infestation, accumulating in plant tissue localised close to aphid feeding after 24 h. There was no clear evidence of systemic plant response to aphid infestation.     

Aphid behavioral responses to virus‐infected plants are similar despite divergent fitness effects

We compared the settling preferences and reproductive potential of an oligophagous herbivore, the pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae), in response to pea plants, Pisum sativum L. cv. ‘Aragorn’ (Fabaceae), infected with two persistently transmitted viruses, Pea enation mosaic virus (PEMV) and Bean leaf roll virus (BLRV), that differ in their distribution within an infected plant. Aphids preferentially oriented toward and settled on plants infected with PEMV or BLRV in comparison with sham‐inoculated plants (plants exposed to herbivory by uninfected aphids), but aphids did not discriminate between plants infected with the two viruses. Analysis of plant volatiles indicated that plants inoculated with either virus had significantly higher green leaf volatile‐to‐monoterpene ratios. Time until reproductive maturity was marginally influenced by plant infection status, with a trend toward earlier nymph production on infected plants. There were consistent age‐specific effects of plant infection status on aphid fecundity: reproduction was significantly enhanced for aphids on BLRV‐infected plants across most time intervals, though mean aphid fecundity did not differ between sham and PEMV‐infected plants. There was no clear pattern of age‐specific survivorship; however, mean aphid lifespan was reduced on plants infected with PEMV. Our results are consistent with predictions of the host manipulation hypothesis, extended to include plant viruses: non‐viruliferous A. pisum preferentially orient to virus‐infected host plants, potentially facilitating pathogen transmission. These studies extend the scope of the host manipulation hypothesis by demonstrating that divergent fitness effects on vectors arise relative to the mode of virus transmission.     

Electrical penetration graphs of the damson-hop aphid, Phorodon humuli on resistant and susceptible hops (Humulus lupulus)

Electrical penetration graphs (EPG's) were used to locate resistance to Phorodon humuli (Schrank) (Homoptera, Aphididae) in hops (Humulus lupulus, Cannabinaceae). Aphids on those hops showing resistance had a much reduced E2 pattern (uptake of phloem). In addition, many aphids on the resistant plants spent time non-probing within two minutes of withdrawing from the phloem. This was not observed with aphids on susceptible hops. The results suggest that resistance is located in the phloem. The involvement of a mechanical factor such as the blocking of aphid stylets, the presence of antinutritional factors, or simply an inadequate supply of nutrients, are discussed as possible resistance mechanisms.     

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.