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.     

Microarray analysis of the interaction between the aphid <Emphasis Type="Italic">Rhopalosiphum padi</Emphasis> and host plants reveals both differences and similarities between susceptible and partially resistant barley lines

The bird cherry-oat aphid (Rhopalosiphum padi L.) is an important pest on cereals causing plant growth reduction without specific leaf symptoms. Breeding of barley (Hordeum vulgare L.) for R. padi resistance shows that there are several resistance genes, reducing aphid growth. To identify candidate sequences for resistance-related genes, we performed microarray analysis of gene expression after aphid infestation in two susceptible and two partially resistant barley genotypes. One of the four lines is a descendant of two of the other genotypes. There were large differences in gene induction between the four lines, indicating substantial variation in response even between closely related genotypes. Genes induced in aphid-infested tissue were mainly related to defence, primary metabolism and signalling. Only 24 genes were induced in all lines, none of them related to oxidative stress or secondary metabolism. Few genes were down-regulated, with none being common to all four lines. There were differences in aphid-induced gene regulation between resistant and susceptible lines. Results from control plants without aphids also revealed differences in constitutive gene expression between the two types of lines. Candidate sequences for induced and constitutive resistance factors have been identified, among them a proteinase inhibitor, a serine/threonine kinase and several thionins. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Phloem specific aphid resistance in Cucumis melo line AR 5: effects on feeding behaviour and performance of Aphis gossypii

The feeding behaviour, excretion rate, and life history traits of the cotton-melon aphid, Aphis gossypii (Glover) (Homoptera, Aphididae), were measured on a resistant melon, Cucumis melo L., breeding line, AR 5. The site of resistance detection by the aphids was determined using the electrical penetration graph (EPG) technique. EPG recordings showed that resistance is expressed within the host plant, rather than on its surface, because the time to first stylet penetration was not significantly different between AR 5 and the closely related susceptible breeding line, PMR 5. EPG patterns associated with stylet pathway activities of the aphids were not significantly different between the resistant and susceptible lines. Significant behavioural differences were observed only after stylets contacted phloem sieve elements. On AR 5, the duration of salivation after sieve element puncture (waveform E1) was significantly longer, and the number of aphids showing phloem sap ingestion (waveform E2) was significantly reduced. We conclude that the resistance mechanism producing the effects seen in this study acts within the phloem sieve elements. Monitoring of excretion rates on the two genotypes showed that aphid feeding was delayed and greatly reduced on the resistant genotype. Comparisons of aphid life history traits and population development between host plant genotypes showed that the effects of resistance act throughout aphid development and are highly effective at slowing down population increase.     

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

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

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.

     

Location of resistance factors in the leaves of potato and wild tuber-bearing Solanum species to the aphid Myzus persicae

Analysis of electrically recorded feeding behaviour of aphids was combined with colony‐development tests to search for sources of resistance to Myzus persicae (Sulzer) (Homoptera: Aphididae) in tuber‐bearing Solanum species (Solanaceae), aiming at a reduction of potato leaf roll virus (PLRV) transmission. Twenty genotypes, originating from 14 gene bank accessions, representing 13 wild tuber‐bearing Solanum spp., three Solanum tuberosum L. (potato) cultivars, and one S. tuberosum breeding line, were selected. Colony‐development tests were carried out in no‐choice experiments by placing adult aphids on plants of each genotype and counting numbers of nymphs and adults on young plants after 8 and 15 days, and on flowering plants after 14 and 30 days. Large differences were observed among genotypes: some developed small colonies and others developed large ones. Also, in a few genotypes, resistance in mature plants was different for leaves of different ages; young leaves were resistant to aphids whereas old senescent leaves were susceptible. The electrical penetration graph (DC‐EPG system) technique was used to study aphid feeding behaviour on each Solanum genotype for 6 h. Electrical penetration graph (EPG) results also showed large differences among the genotypes, indicating resistance at the leaf surface and at three different levels of plant tissue (epidermis, mesophyll, and phloem). Therefore, it was concluded that different mechanisms of resistance to M. persicae exist among the genotypes analysed. EPGs recorded from aphids on Solanum berthaultii Hawkes and Solanum tarijense Hawkes with and without glandular trichomes showed that strong surface resistance can bias EPG parameters associated with resistance located in deeper tissues. Experimental evidence is presented that the resistance to aphids in the genotypes with glandular trichomes strongly depends on these morphological structures.     

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.     

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.     

Russian wheat aphid biotype RWASA2 causes more vascular disruption than RWASA1 on resistant barley lines

We investigated the comparative effects of the feeding damage caused by two Russian wheat aphid (RWA, Diuraphis noxia Kurdjumov) biotypes, RWASA1 and RWASA2, on leaves of three RWA-resistant barley (Hordeum vulgare L.) lines from the USDA-ARS, and used a South African non-resistant cultivar as control. The relationship between aphid breeding capacity and the structural damage inflicted by the aphids was studied, using wide-field fluorescence and transmission electron microscopy (TEM). Colonies of the two biotypes grew rapidly on all four barley lines during a 10 day feeding exposure but as expected, population size and density were generally lower on the resistant lines than on the non-resistant cultivar. The new South African biotype, RWASA2, bred significantly faster than the original RWASA1 biotype. The feeding and water uptake-related damage sustained by phloem and xylem tissues of the resistant lines suggest that RWASA2 was a more aggressive feeder and caused substantially more cell damage than RWASA1. Examination of wound callose distribution after aphid feeding revealed that high levels of wound callose occurred in non-resistant and in resistant lines. Reduction in aphid population size, as well as ultrastructural damage during feeding by RWA biotypes on resistant lines, signals potential antibiotic and tolerant responses of the barley lines to aphid feeding. We infer from callose distribution and ultrastructural studies, that phloem transport would be substantially reduced in the non-resistant PUMA and to a lesser extent in the resistant STARS lines, which suggests that the STARS lines may be a potential source of RWASA1 and RWASA2-resistance.     

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.     

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

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

Diurnal feeding as a potential mechanism of osmoregulation in aphids

Diurnal variation in phloem sap composition has a strong infuence on aphid performance.The sugar-rich phloem sap serves as the sole diet for aphids and a suite of physiological mechanisms and behaviors allowv them to tolerate the high osmotic stress.Here,we tested the hypothesis that night-time feeding by aphids is a behavior that takes advantage of the low sugar diet in the night to compensate for osmotic stress incurred while feeding on high sugar diet during the day.Using the electrical penetration graph(EPG)technique.we examined the eiects of diurmal rhythm on feeding behaviors of bird cherry-oat aphid(Rhopalosiphurm padi L.)on wheat.A strong diurmal rhythm in aphids as indicated by the presence of a cyclical pattern of expression in a core clock gene did not impact aphid feeding and similar feeding behaviors were observed during day and night.The major difference observed between day and night feeding was that aphids spent significantly longer time in phloem salivation during the night compared to the day.In contrast,aphid hydration was reduced at the end of the day-time feeding compared to end of the night-time fepding.Gene expression analysis of R.padi osmoregulatory genes indicated that sugar break down and water transport into the aphid gut was reduced at night.These data suggest that while diumal variation occurs in phloem sap composition,aphids use night time feeding to overcome the high osmotic stress incurred while feeding on sugar-rich phloem sap during the day.     

Development and reproductive potential of the cereal aphid Sitobion avenae on resistant wheat lines (Triticum monococcum)

The effect of wheat resistance in lines of Triticum monococcum L., on the reproductive performances of the cereal aphid (Sitobion avenae F.) was investigated. Aphids were reared from birth to adult moult either on resistant or susceptible wheat lines, and transferred as apterae to both host genotypes. The influence of these transfers on the subsequent adult weight, gonad status and reproductive performances was evaluated. Aphids transferred from resistant to susceptible plants proved able to compensate for their poor nymphal growth, mainly through additional embryo growth and an increase in the number of matured embryos within the first 10 days of their adult life. Most aphids transferred from susceptible to resistant plants died within the first week following the transfer. Their most advanced embryos matured and were born, but subsequent embryo growth was quickly reduced. The reproductive strategies adopted by S. avenae when facing plant resistance, and the hypothesis of a resistant mechanism based on a poor nutritional state of the resistant plants are discussed.     

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.     

Possible roles of esterase, glutathione S-transferase, and superoxide dismutase activities in understanding aphid–cereal interactions

Activities of the detoxification enzymes esterase, glutathione S‐transferase, and of superoxide dismutase in aphids and aphid‐infested cereal leaves were assayed using polyacrylamide gel electrophoresis and a spectrophotometer to elucidate the enzymatic mechanisms of aphid resistance in cereal plants. A chlorosis‐eliciting Russian wheat aphid, Diuraphis noxia (Mordvilko), and non‐chlorosis‐eliciting bird cherry‐oat aphid, Rhopalosiphum padi (L.), and four cereals were used in this study. The four cereal genotypes were ‘Arapahoe’ (susceptible) and ‘Halt’ (resistant) wheat (Triticum aestivum L.), ‘Morex’ (susceptible) barley (Hordeum vulgare L.), and ‘Border’ (resistant) oat (Avena sativa L.). Esterase isozymes differed between the two aphid species, although glutathione S‐transferase and superoxide dismutase did not. Esterase, glutathione S‐transferase, and superoxide dismutase activities in either aphid species were not affected by the level of resistance of a cereal to D. noxia. The assays of cereal leaf samples showed that D. noxia feeding elicited an increase in esterase activity in all four cereal genotypes, although R. padi feeding did not. The increase of esterase activity in cereals, however, was not correlated to aphid resistance in the cereals. The time‐series assays of aphid‐infested cereal leaves showed that D. noxia‐infested Morex barley had a significant increase in esterase activity on all sampling dates (3, 6, and 9 days) in comparison with either uninfested or R. padi‐infested barley. No difference in glutathione S‐transferase activity was detected among either aphid infestations or sampling dates. The electrophoretic assays, however, revealed that aphid feeding elicited a significant increase in superoxide dismutase activity, which served as the control of glutathione S‐transferase activity assays. The increase in esterase and superoxide dismutase activities suggested that D. noxia feeding imposes not only toxic, but also oxidative stresses on the cereals. The ramification of using these enzyme activity data to understand the etiology of D. noxia‐elicited chlorosis is discussed.     

Role of phloem sap quality and exudation characteristics on performance of pea aphid grown on lucerne genotypes

The phloem sap of two lucerne (Medicago sativa L.) genotypes inducing differences in pea aphid,Acyrthosiphon pisum Harris (Homoptera: Aphididae) performances (weight, fecundity, survival), was collected by stylectomy (radio frequency microcautery). Sugars and amino acids were assayed. No significant difference in their concentrations could be established between genotypes. It is thus unlikely that a substantial part of the varietal resistance of these lucerne genotypes to pea aphid could be related to the content in sugars or amino acids. The same conclusion was drawn for the sugar/amino acid ratio. The amino acid profiles were globally similar. They nevertheless allowed a discrimination between the two genotypes on a few amino acids (alanine, leucine, isoleucine, arginine, ornithine) only. Moreover, the degree of resistance can be related neither to GABA concentration, nor to that of sulfur amino acids, because these constituants were either absent, or in trace amounts, whatever the lucerne genotype taken into account. Stylectomy parameters revealed some interesting differences in the sap exudation of the two genotypes. Average volume of exudation by stylet and average duration of exudation were respective 2.8 and 2.3 times higher in the susceptible genotype. These results suggest a reduced flow of phloem sap, after puncturing of the sieve-tubes by the aphid stylets on the resistant genotype; this could result in a greater difficulty in phloem sap ingestion by aphids settled on the resistant genotype and thus in suboptimal performances.     

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.     

Population growth rate and relative virulence of the two South African biotypes of Russian wheat aphid,Diuraphis noxia,and bird cherry‐oat aphid,Rhopalosiphum padi,on resistant and non‐resistant barley

In South Africa a new biotype of the Russian wheat aphid (RWA), Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae), RWASA2, has appeared which exhibits an improved performance compared to the original biotype (RWASA1) on wheat containing the Dn1 resistance gene. We examined population growth rates as well as damage caused by RWASA1 and RWASA2, in addition to a different aphid species, the bird cherry‐oat aphid (BCA), Rhopalosiphum padi L. (Hemiptera: Aphididae), on three RWA‐resistant barley [Hordeum vulgare L. (Poaceae)] lines (STARS‐9577B, STARS‐0502B, and STARS‐9301B) and one susceptible control (PUMA). RWASA2 had a higher reproductive rate than RWASA1 on all barley lines tested, which is consistent with previous results on wheat. Two of the RWA‐resistant lines (STARS‐0502B and STARS‐9301B) also exhibited a similar resistance phenotype against BCA. In our experiments, severe chlorosis and leaf roll appeared earlier on the control PUMA barley variety as a result of RWASA2 feeding than was the case with RWASA1, probably due to the differences in reproductive rate. Although chlorosis appeared earlier on resistant plants after RWASA2 feeding, this symptom developed much faster during RWASA1 feeding on all three resistant lines tested. As chlorosis did not correlate well with aphid population numbers, we surmise that the differential chlorosis effects may be related to differences in the amount of saliva introduced by the two aphid clones during feeding. Our results indicate that the difference between RWASA2 and RWASA1 are broader than a ‘gene for gene’ interaction with the Dn1 resistance (R) gene in wheat, and that these biotypes also differ in important aspects of their biology.     

Genotypic variation in induced resistance and induced susceptibility in the peach-Myzus persicae aphid system

In response to herbivore damage or stress, plants may express physiological or morphological changes known as induced responses. We tested whether previous herbivory by the aphid Myzus persicae differentially altered the expression of resistance and susceptibility among five genotypes of peach that differ in their resistance phenotype (avoidance resistance, antibiosis resistance or susceptibility). We measured behavioural and performance parameters of aphid success on plants previously infested by conspecifics as compared to uninfested controls. Significant variation was found both among genotypes and among resistance phenotype, including between genotypes showing a same resistance phenotype. Genotypes with avoidance resistance showed either induced resistance to aphid settling or no response. Genotypes with antibiosis resistance showed induced susceptibility to aphid settling, but the effects of previous herbivory on aphid development were either positive or negative depending on the genotype. In the susceptible genotype, most parameters of aphid settlement and performance, including reproduction, were positively influenced by previous herbivory. Using electronic recording, the aphid probing behaviour was examined to tentatively identify host plant tissues most likely to play a role in induced defenses. Probing behaviour was significantly affected by plant genotype, previous herbivory, and their interaction, indicating complex relations between the two factors. In the genotypes with avoidance resistance, aphids were deterred before they reach the phloem. In the genotypes expressing susceptibility or antibiosis resistance, previous herbivory triggered instead the induction of a phloem‐mediated response, that however diverged depending on the resistance status (facilitation or reduction of phloem sap uptake respectively). Genotypic variation in induction found in the peach‐M. persicae system establishes a useful framework to improve our knowledge of the ecological role of induced plant responses to aphids.