Feeding behavior and performance of Nasonovia ribisnigri on grafts,detached leaves,and leaf disks of resistant and susceptible lettuce

Aphids are dependent on the phloem sap of plants as their only source of nutrients. Host‐plant resistance in lettuce, Lactuca sativa L. (Asteraceae), mediated by the Nr gene is used to control the lettuce aphid Nasonovia ribisnigri (Mosely) (Hemiptera: Aphididae). The resistance is located in the phloem; however, the exact mechanism of resistance is unknown. In this study, we investigated whether the resistance factor (or factors) is synthesized in the root or in the shoot. The feeding behavior and performance of avirulent N. ribisnigri were studied on grafts of resistant and susceptible lettuce. In addition, the persistence of resistance in excised lettuce tissue was measured, by studying the feeding behavior and performance of N. ribisnigri on detached leaves and leaf disks of resistant lettuce. It appears that the resistance factor encoded by the Nr gene is produced in the shoots: aphid feeding was reduced on resistant shoots grafted on susceptible roots, whereas aphids were able to feed on grafts of susceptible shoots on resistant roots. Partial loss of resistance was observed after detachment of leaves and excision of leaf disks from resistant plants. Aphids fed longer on excised resistant plant tissue compared with intact resistant plants; however, compared with excised plant tissue of the susceptible cultivar, the time spent on feeding was shorter, indicating resistance was not completely lost. Our findings caution against the use of excised leaf material for aphid resistance bioassays.     

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.     

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.     

The impact of Meu1-mediated resistance in tomato on longevity, fecundity and behavior of the potato aphid, Macrosiphum euphorbiae

The effect of the tomato resistance gene, Meu1, on feeding, longevity, fecundity and developmental rate of the pink biotype of the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera, Aphididae), was determined using nearly isogenic tomato (Lycopersicon esculentum Mill, Solanaceae) lines. Aphid mortality was significantly higher on resistant plants, with 60% of the aphids dying by the 4th day of exposure. By the 10th day, all the aphids on the resistant plants were dead whereas 100% of the aphids on susceptible plants were alive. Meu1-mediated resistance resulted in significantly decreased fecundity with a ten-fold decrease in the net fertility rate (4.5 and 45.7 progeny per aphid on resistant and susceptible tomato, respectively). A qualitative analysis showed that honeydew was produced by aphids on resistant and susceptible plants, suggesting that aphids initiate feeding on both plant types. However, significantly lower quantities of honeydew were present when aphids were caged on resistant plants. There were also significant differences in aphid location on resistant and susceptible leaves. Experiments evaluating behavior in less than 24 h showed that aphids left resistant leaves after relatively short exposure (3–6 h). Aphids transferred from resistant to susceptible tomato at intervals between 3 h and 24 h resumed feeding as evidenced by presence of honeydew. Although the mechanism by which Meu1-mediated resistance operates is not yet known, our data suggest that resistance factors act rapidly after initiation of feeding and that lower fecundity and longevity are related to reduction in aphid feeding.     

The effect of leaf excision on the performance of Myzus persicae and Brevicoryne brassicae in relation to the nutrient treatment of the plants

Two groups of brussels sprout plants of one variety given different fertilizer treatments proved ‘resistant’ and ‘susceptible’ to both Myzus persicae and Breicoryrie brassicae, as measured by aphid mean relative growth rate. The hypothesis that leaf disc would fail to reflect differences in ‘nutritionallybased’ plant resistance was confirmed with B.brussicue but not M.persicae. Both aphids grew more slowly on discs than leaves of ‘susceptible’ plants; the reverse tendency showed on ‘resistant’ material. Total:(, amino nitrogen proved poorly correlated with aphid performance. Previously published (van Emden & Bashford, 1971) equations relating individual amino acids to aphid performance gave a better fit to the results, but still failed to account for the reduced performance of the aphids on ‘susceptible’ discs.     

Comparison of feeding behaviour of two Brassica pests Brevicoryne brassicae and Myzus persicae on wild and cultivated brassica species

Feeding behaviour of the specialist Brassicae aphid, Brevicoryne brassicae (L) (cabbage aphid) and the generalist, Myzus persicae, (Sulzer) (peach potato aphid) was monitored electronically on the susceptible cauliflower, Brassica oleracea var. botrytis cv Newton Seale, and a range of 17 Brassica species, B. carinata, B. juncea, B. nigra, B. macrocarpa, and B. villosa var. drepanensis and cultivated brassica varieties, B. oleracea, B. campestris and B. napus. Aphids, monitored for 10 h on the underside of leaves, performed recognisable feeding behaviour on all brassica species. The main differences in feeding behaviour, between M. persicae and B. brassicae, on the susceptible cauliflower Newton Seale, were fewer probes, shorter times to initially reach the phloem but longer times to establish sustained phloem ingestion and the longer times spent, by M. persicae, in xylem ingestion.Feeding behaviour on the range of brassica species tested indicated that generalist and specialist aphids are influenced differently by the host plant. A longer time spent in xylem ingestion was again the major difference in the feeding behaviour of the two aphids. In addition, rejection of passive phloem ingestion, by M. persicae, was not related so closely to increased time spent in non probing activities, as for B. brassicae. This observation indicates that M. persicae does not generally accept or reject brassica species due to the presence of phagostimulants, such as glucosinolates at the leaf surface or along the stylet pathway, unless the concentration is very high. Differences in feeding strategies employed by generalist and specialist aphids on the same plants are discussed.     

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.     

高粱抗高粱蚜的生化基础

高粱中的可溶性总氮、可溶性总糖和绝大部分游离氨基酸,特别是必需氨基酸,在感性品种中的含量显著地比抗性品种中的含量高.感性品种叶液微偏酸,抗性品种近中性.多元回归分析表明,高梁对高粱蚜Melanaphis sacchari(Zehntner)的抗性与可溶性总氮、可溶性总糖和游离氨基酸的含量呈反相关.     

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.

     

Effect of bioregulator-treated sorghum on greenbug fecundity and feeding behavior: implications for host-plant resistance

Three commercial and six experimental plant growth bioregulators were surveyed for their effect on aphid reproduction when applied to sorghum. Only CCC and PIX had a significant effect on the greenbug, Schizaphis graminum. Application of the commercial bioregulators CCC and PIX^R caused about a 50% decrease in aphid reproduction rate when applied to greenbug susceptible sorghum but had little effect when applied to a greenbug resistant sorghum line. Electronic monitoring of aphid probing behavior on CCC treated, greenbug-susceptible sorghum showed a response pattern which was indistinguishable from that normally observed on greenbug resistant lines and was different from that associated with aphid probing behavior on untreated susceptible lines. The isolated pectin content of the CCC treated susceptible sorghum was twice that of the controls and had twice the methoxy content. These results support the argument that pectin is a barrier to aphid-stylet penetration for phloem feeding aphids which probe intercellularly and that manipulation of pectin content and/or structure can be a major factor in host-plant resistance to sap-sucking insects.     

Analysis of wheat resistance to the cereal aphidSitobion avenae using electrical penetration graphs and flow charts combined with correspondence analysis

The behaviour ofSitobion avenae (F.), was compared on resistant wheat lines ofTriticum monococcum (L.) and a susceptible variety ofTriticum aestivum (L.). Firstly, stylet penetration activities were monitored with the Electrical Penetration Graph (EPG) technique and subsequently analysed using flow charts combined with correspondence analysis. Plant resistance was shown to be associated with repeated penetrations without access to either the xylem or the phloem, and with numerous failures in starting a sustained sap ingestion (as represented by pattern E2). Access to sieve elements of the phloem did not seem to be much affected on resistant plants but it took the aphid three times as long to produce a sap ingestion pattern when maintained on the resistant lineT. monococcum n^o 44 (Tm44) as compared with aphids maintained on susceptible plants. As a result the total time spent in ingesting from sieve elements was reduced by 72% on Tm44. Secondly, direct observations of freely-moving apterous adults were performed. Aphids did not discriminate between resistant and susceptible wheat during the first 30 min of access to test leaves, but only 4 out of 25 aphids were still probing after eight hours on resistant Tm44. The relevance of these results to possible location of the resistance factor(s) are discussed. Although detection of plant resistance before sieve elements are reached can not be rigorously excluded, the factors involved inT. monococcum resistance toS. avenae undoubtedly occur within the phloem vessels.     

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.     

Locating a resistance mechanism to the cabbage aphid in two wild Brassicas

Feeding behaviour of the cabbage aphid,Brevicoryne brassicae, was monitored electronically on two resistantBrassica species,B. fruticulosa andB. spinescens, and compared with a susceptible controlB. oleracea var.capitata cv. Offenham Compacta. Aphids, monitored for 10 h on the under side of leaves, performed recognizable feeding behaviour on all species. Electrical Penetration Graphs (EPGs) of aphids on resistant and susceptible plants showed no difference in behaviour for aphids on resistantBrassica species compared to susceptible until stylets penetrated the phloem sieve elements when a large reduction in the duration of passive phloem uptake (E₂ pattern) onB. fruticulosa was indicated. Although feeding behaviour on 6 week-old plants ofB. spinescens was similar to the susceptible controls, behaviour on 10 week-old plants was similar to that recorded forB. fruticulosa. The mechanism of resistance is thought to be located in the sieve element as the normal sieve element salivation (E₁) signal was either quickly terminated by withdrawal of the stylets from the sieve element or continued as a disrupted E₂ pattern. Analysis of secondary plant compounds in the threeBrassica species only identified significant differences in the glucosinolate profile. No reproducible differences were detected in the concentration of phenolics or anthocyanins. The major glucosinolate component ofB. fruticulosa andB. spinescens was gluconapin rather than glucobrassicin and glucoiberin as found in the susceptible host plant. However, both pure glucosinolates and glucosinolate extracts from all three species did not reduce aphid survival on chemically-defined artificial diets. These results suggest that the mechanism of resistance may be a mechanical blocking of the sieve element or stylets rather than a difference in the secondary plant chemistry of glucosinolates and phenolics.     

Probing behaviour of the green peach aphid Myzus persicae on resistant Prunus genotypes

Electrical penetration graphs of Myzus persicae (Sulzer) (Homoptera: Aphididae) feeding behaviour on four resistant and two susceptible genotypes of peach (Prunus persica L. Batsch) and related species showed that resistance was mainly linked to (i) reduced duration of phloem sap uptake, (ii) reduced percentage of pattern E1 (salivary secretion into sieve elements) followed by pattern E2 (sap ingestion) and (iii) increased number of shifts from E1 to E2 and back. These results suggest the unsuitability of phloem sap, and thus repetitive failures to initiate sustained ingestion. Extensive comparisons of the EPGs also revealed more specific trends. Aphids on the most susceptible cultivar GF305 produced significantly longer potential drops than on other peach genotypes. On the resistant Rubira, aphids generated more penetrations before the first E occurred, indicating the possible presence of a resistance factor before the phloem was reached. The clone P1908 of the wild species Prunus davidiana displayed traits of both susceptibility (less but longer probes) and resistance. In particular, aphids produced more E1, suggesting difficulties in preparing sieve elements before feeding. The aphid probing process could be correlated with aphid settling behaviour and bionomics, as previously reported, and gave evidence for the existence of different mechanisms underlying resistance in the tested genotypes against M. persicae.     

Feeding behaviour of Aphis craccivora (Koch) on cowpea cultivars with different levels of aphid resistance

The feeding activity of Aphis craccivora (Koch) was monitored on cowpea plants of aphid-susceptible (Vita 7) and aphid-resistant (TVu 801) cultivars, using an AC electronic feeding monitoring system. Waveforms corresponding to salivation, phloem ingestion and non-phloem ingestion were observed. Aphids probing on the resistant cultivar showed a significantly reduced ingestion of phloem sap compared with those aphids which fed on the susceptible cultivar. The insect also made more brief and repeated probes on TVu 801. In addition, the duration of non-probing activities and non-phloem ingestion was shorter on the susceptible than on the resistant cultivar.     

How nutritionally imbalanced is phloem sap for aphids?

Aphids harbour intracellular symbionts (Buchnera) that provide their host with amino acids present in low amounts in their diet, phloem sap. To find out the extent to which aphids depend on their symbionts for synthesis of individual essential amino acids, we have evaluated how well phloem sap amino acid composition matches the aphids' needs. Amino acid compositions of the ingested phloem sap were compared to amino acids in aphid body proteins and also to available information about optimal diet composition for other plant feeding insects. Phloem sap data from severed stylets of two aphid species, Rhopalosiphum padi (L.) (Homoptera: Aphididae) feeding on wheat, and Uroleucon sonchi (L.) (Homoptera: Aphididae) feeding on Sonchus oleraceus (L.), together with published information on phloem sap compositions from other plant species were used.Phloem sap has in general only around 20% essential amino acids, with a range from 15–48%. Aphid body proteins and optimal diets for two other plant feeding insects have around 50%. The phloem sap of early flowering S. oleraceus ingested by U. sonchi contained 48%, which seems to be exceptional. Aphids feeding on different plants appear to be very differently dependent on their symbionts for their overall essential amino acid synthesis, due to the large variation in proportion of essential amino acids in phloem sap from different plants.The profile of the essential amino acids in phloem sap from different plant species corresponds rather well to profiles of both aphid body proteins and optimal diets determined for plant feeding insects. However, methionine and leucine in phloem sap are in general low in these comparisons, suggesting a higher dependence on the symbiont for synthesis of these amino acids. Concentrations of several essential amino acids in phloem from different plant species seem to vary together, suggesting that levels of symbiont provisioning of different amino acids are adjusted in parallel.     

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.     

植物和刺吸式口器昆虫的诱导防御与反防御研究进展

刺吸式口器昆虫在长期的进化过程中形成特殊的口针结构,用于专门吸食植物韧皮部筛管细胞的汁液成分.以蚜虫为例,它们在取食过程中分泌的胶状唾液和水状唾液将有效的降低植物防御反应,其中水状唾液包含的大量酶类不仅可以帮助蚜虫穿刺植物韧皮部,刺探到筛管细胞,同时也是植物感受蚜虫为害的激发因子,诱导出植物防御反应和相关抗性基因的表达...     

Location of the mechanism of resistance to Amphorophora agathonica (Hemiptera: Aphididae) in red raspberry

The aphid Amphorophora agathonica Hottes (Hemiptera: Aphididae) is an important virus vector in red (Rubus idaeus L.) and black (Rubus occidentalis L.) raspberries in North America. Raspberry resistance to A. agathonica in the form of a single dominant gene named Ag1 has been relied upon to help control aphid-transmitted plant viruses; however, the mechanism of resistance to the insect is poorly understood. Aphid feeding was monitored using an electrical penetration graph on the resistant red raspberry 'Tulameen' and compared with a susceptible control, 'Vintage'. There were no differences in pathway feeding behaviors of aphids as they moved toward the phloem. Once in the phloem, however, aphids feeding on resistant plants spent significantly more time salivating than on susceptible plants, and ingested significantly less phloem sap. This suggests that a mechanism for resistance to A. agathonica is located in the phloem. Reduced ingestion of phloem may result in inefficient acquisition of viruses and is a likely explanation for the lack of aphid-transmitted viruses in plantings of resistant cultivars.     

Feeding damage by Diuraphis noxia results in a nutritionally enhanced phloem diet

The feeding behavior of Diuraphis noxia Mordvilko (Homoptera: Aphididae) on susceptible hosts causes both ultrastructural and tissue level damage which may affect phloem composition. Genetic evidence suggests that endosymbiotic bacteria in most aphids overproduce limiting amino acids to benefit hosts but that D. noxia depends less on endosymbionts for these nutrients, possibly due to an enriched diet. To determine whether D. noxia feeding damage results in higher concentrations of essential amino acids, stylet exudates were analyzed from wheat (Triticum aestivum) damaged to different degrees. Comparison of samples from undamaged and damaged susceptible wheat revealed changes in amino acid composition and an increase in levels of essential amino acids, indicating a nutritionally enhanced ingesta. The changes in stylet exudates paralleled changes in leaf exudates, indicating that the effects are systemic. Feeding damage is not observed on a resistant wheat host, var. Halt, and leaf exudates from infested Halt did not show changes in amino acid composition. Mean relative growth of nymphs was significantly lower on Halt than on susceptible Arapahoe, indicating that Halt is a less suitable host. Both varieties show similar amino acid levels in non-infested samples, suggesting that D. noxia infestation does not enhance the phloem environment in Halt. This study provides evidence that aphid feeding can generate a nutritionally enhanced phloem diet.