Characterization of pea aphid resistance in Medicago truncatula

To achieve a thorough understanding of plant-aphid interactions, it is necessary to investigate in detail both the plant and insect side of the interaction. The pea aphid (PA; Acyrthosiphon pisum) has been selected by an international consortium as the model species for genetics and genomics studies, and the model legume Medicago truncatula is a host of this aphid. In this study, we identified resistance to PA in a M. truncatula line, 'Jester', with well-characterized resistance to a closely related aphid, the bluegreen aphid (BGA; Acyrthosiphon kondoi). The biology of resistance to the two aphid species shared similarity, with resistance in both cases occurring at the level of the phloem, requiring an intact plant and involving a combination of antixenosis, antibiosis, and plant tolerance. In addition, PA resistance cosegregated in 'Jester' with a single dominant gene for BGA resistance. These results raised the possibility that both resistances may be mediated by the same mechanism. This was not supported by the results of gene induction studies, and resistance induced by BGA had no effect on PA feeding. Moreover, different genetic backgrounds containing a BGA resistance gene from the same resistance donor differ in resistance to PA. These results suggest that distinct mechanisms are involved in resistance to these two aphid species. Resistance to PA and BGA in the same genetic background in M. truncatula makes this plant an attractive model for the study of both plant and aphid components of resistant and susceptible plant-aphid interactions.     

Independent action and contrasting phenotypes of resistance genes against spotted alfalfa aphid and bluegreen aphid in Medicago truncatula

Host resistance to aphids is poorly understood. Medicago truncatula, a model legume and cultivated pasture species, was used to elucidate defense against two aphid species, Therioaphis trifolii f. maculata (spotted alfalfa aphid, SAA) and Acyrthosiphon kondoi (bluegreen aphid, BGA). Aphid performance and plant damage were compared between near-isogenic cultivars, Mogul and Borung, that differ in resistance to both aphids. Analyses of aphid resistance in Mogul x Borung F2 plants and their progeny revealed modes of action and chromosome locations of resistance genes. Separate genes were identified for SAA resistance (TTR) and BGA resistance (AKR); both mapped to chromosome 3 but were found to act independently to reduce survival and growth of their target aphid species. The TTR locus controls distinct, and contrasting, local and systemic plant responses between the near-isogenic cultivars. TTR-mediated plant responses imply interaction between a resistance factor(s) in vascular tissue and a bioactive component(s) of SAA saliva. Features of both resistance traits suggest homology to aphid resistance in other legumes; elucidation of their molecular mechanisms will likely apply to other aphid-plant interactions.     

Tolerance of antibiotic and susceptible cereal seedlings to the aphids Metopolophium dirhodum and Rhopalosiphum padi

Some cereal seedlings exhibit antibiotic and antixenotic resistance to the aphids Metopolophium dirhodum (Walker) and Rhopalosiphum padi (L.), because the seedlings contain hydroxamic acids or gramine. The association between tolerance to aphids and aphid antibiosis was investigated for three cereals, Dollarbird wheat Vulcan wheat and Yagan barley. The dry biomass gained by the aphids and the simultaneous reduction in the biomass of the plants (biomass conversion ratio) quantified tolerance. Biomass production and the density dependence of biomass production by the aphids quantified antibiosis more effectively than fecundity. Vulcan wheat, which has more hydroxamic acid than Dollarbird wheat showed the highest level of antibiosis, and the barley was not antibiotic for either aphid. The biomass conversion ratio was a constant; the biomass of an infested plant was reduced by 3 mg for each mg of aphid biomass gained, regardless of aphid species, plant cultivar, or aphid density. The three plants showed no differential tolerance to the aphids, and therefore tolerance is not associated with antibiosis in this case.     

Categories of Resistance to Greenbug and Yellow Sugarcane Aphid (Hemiptera: Aphididae) in Three Tetraploid Switchgrass Populations

Switchgrass, Panicum virgatum L., has been targeted as a bioenergy feedstock. However, little is currently known of the mechanisms of insect resistance in this species. Here, two no-choice studies were performed to determine the categories (antibiosis and tolerance) and relative levels of resistance of three switchgrass populations (Kanlow–lowland ecotype, Summer–upland ecotype, and third generation derivatives between Kanlow?×?Summer plants, K×S) previously identified with differential levels of resistance to the greenbug, Schizaphis graminum (Rondani), and yellow sugarcane aphid, Sipha flava (Forbes). No-choice studies indicated that Kanlow possessed multi-species resistance, with high levels of antibiosis to both aphid species, based on aphid survival at 7 and 14 days after aphid introduction and cumulative aphid days, while K×S possessed low-to-moderate levels of antibiosis to S. flava. Further, functional plant loss indices based on plant height and biomass indicated that tolerance is an important category of resistance for Summer plants to S. graminum. These studies also indicated that Summer lacks both tolerance and antibiosis to S. flava, relative to the other switchgrasses tested, whereas K×S lack tolerance and antibiosis to S. graminum. These studies are the first attempt to analyze the categories of resistance in switchgrass and provide critical information for characterizing the biological mechanisms of resistance and improving our knowledge of the plant–insect interactions within this system.     

Seedling and adult plant resistance to Sitobion avenae (Hemiptera: Aphididae) in Triticum monococcum (Poaceae), an ancestor of wheat

Cereal aphids are important pests of wheat, Triticum aestivum L. and Triticum durum Desf. Crop resistance is a desirable method for managing cereal aphids in central North America, where the dominant crop, spring-sown wheat, has a low value per unit area. A diploid ancestor of wheat, Triticum monococcum L., is reported to be partially resistant to Sitobion avenae (Fabricius), the most damaging cereal aphid in the region. To identify potential sources of resistance, 42 accessions of T. monococcum and three cultivated wheats were infested with aphids, seedlings for six days and adult plants for 21 days. Overall resistance was estimated by the biomass loss of foliage and spikes in relation to uninfested control plants. Antibiosis was estimated by the gain in biomass of aphids during infestation, and tolerance was estimated as a biomass conversion ratio, overall resistance divided by antibiosis. A few T. monococcum accessions exhibited partial resistance. No relationship was found between seedling and adult plant resistance: the former exhibited primarily antibiosis and the latter primarily tolerance. Two accessions with antibiosis reduced aphid biomass by 60% compared with commercial wheats. Tolerance was correlated with growth potential, and was useful only in accessions with high growth potential. Four accessions exhibited tolerance levels at least 30% greater than commercial wheats. Highly susceptible accessions also were identified, which would be useful for investigating the inheritance of antibiosis and tolerance.     

Patterns of resistance to three cereal aphids among wheats in the genus Triticum (Poaceae)

Forty-one accessions of wild and cultivated wheats belonging to 19 Triticum species were tested in the field for resistance to three species of aphids, Rhopalosiphum padi Linnaeus, Sitobion avenae Fabricius and Schizaphis graminum Rondani. Antibiotic resistance was estimated by the increase in biomass of aphids over 21 days on adult plants. Overall resistance was estimated by the plant biomass lost due to aphid infestation. All three species of aphids survived and reproduced on all wheats, and reduced spike biomass compared to uninfested controls. The level of antibiosis varied among wheat species and among accessions, with accessions from three, five and one species showing antibiosis to R. padi, S. avenae and S. graminum, respectively. Overall resistance to the three aphid species was observed in five to seven accessions per aphid species. Resistance was usually specific to one aphid species. The frequency of accessions with antibiosis or overall resistance was associated with the ploidy level of the plant species. Except for overall resistance to R. padi, resistance was highest for diploid species and lowest for hexaploid species. No consistent relationship between resistance and level of domestication was detected. Accessions of the wild wheats, Triticum boeoticum Bois, Triticum tauschii (Coss.) Schmal. and Triticum araraticum Jakubz. exhibited high levels of resistance to aphids, as did Triticum monococcum L. which is derived from T. boeoticum. Nevertheless, individual susceptible or resistant accessions occurred at all levels within the evolutionary tree of wheat.     

Characterization of the interaction between the bacterial wilt pathogen Ralstonia solanacearum and the model legume plant Medicago truncatula

The soilborne pathogen Ralstonia solanacearum is the causal agent of bacterial wilt and attacks more than 200 plant species, including some legumes and the model legume plant Medicago truncatula. We have demonstrated that M. truncatula accessions Jemalong A17 and F83005.5 are susceptible to R. solanacearum and, by screening 28 R. solanacearum strains on the two M. truncatula lines, differential interactions were identified. R. solanacearum GMI1000 infected Jemalong A17 line, and disease symptoms were dependent upon functional hrp genes. An in vitro root inoculation method was employed to demonstrate that R. solanacearum colonized M. truncatula via the xylem and intercellular spaces. R. solanacearum multiplication was restricted by a factor greater than 1 x 10(5) in the resistant line F83005.5 compared with susceptible Jemalong A17. Genetic analysis of recombinant inbred lines from a cross between Jemalong A17 and F83005.5 revealed the presence of major quantitative trait loci for bacterial wilt resistance located on chromosome 5. The results indicate that the root pathosystem for M. truncatula will provide useful traits for molecular analyses of disease and resistance in this model plant species.     

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.     

Characterization of resistance to multiple aphid species (Hemiptera: Aphididae) in Medicago truncatula

Aphids are phloem-feeding insects that damage many important crops throughout the world yet, compared to plant-pathogen interactions, little is known about the mechanisms by which plants become resistant to aphids. Medicago truncatula (barrel medic) is widely considered as the pre-eminent model legume for genetic and biological research and in Australia is an important pasture species. Six cultivars of M. truncatula with varying levels of resistance to two pests of pasture and forage legumes, the bluegreen aphid Acyrthosiphon kondoi Shinji and the spotted alfalfa aphid Therioaphis trifolii f. maculata. (Buckton) are investigated. Two resistance phenotypes against T. trifolii f. maculata are described, one of which is particularly effective, killing most aphids within 24 h of infestation. Each resistance phenotype provided a similar but somewhat less effective degree of resistance to the closely-related spotted clover aphid Therioaphis trifolii (Monell). In the case of A. kondoi only one resistance phenotype was observed, which did not vary among different genetic backgrounds. None of the observed resistance against A. kondoi or T. trifolii f. maculata significantly affected the performance of green peach aphid Myzus persicae (Sulzer) or cowpea aphid Aphis craccivora Koch. The existence of multiple aphid resistance mechanisms in similar genetic backgrounds of this model plant provides a unique opportunity to characterize the fundamental basis of plant defence to these serious agricultural pests.     

Plant resistance components of two greenbug (Homoptera: Aphididae) resistant wheats

Several biotypes of the greenbug, Schizaphis graminum (Rondani), attack winter wheat, Triticum aestivum L., on the Southern Plains every year. Two wheat germplasm sources of resistance ('Largo' and 'GRS 1201') have been developed that provide protection against the three predominant greenbug biotypes (E, I, and K). Each source has agronomic and end-use quality advantages and disadvantages for the breeder to consider in choosing a greenbug-resistant breeding line. We compared these two germplasms to determine their levels of resistance against biotype E. Components of resistance (i.e., antibiosis, antixenosis, and tolerance) were measured on seedlings of GRS 1201, Largo, and 'TAM W-101' (a susceptible control). Several aphid and plant measurements (e.g., total number of aphids produced per plant, aphid selection preferences, and plant damage ratings) were recorded for each plant entry. Select data recorded for each resistance component were normalized and combined to derive a plant resistance index for each wheat entry. Results indicated that GRS 1201 had a higher level of combined resistance components than did Largo, followed by TAM W-101, the susceptible control. These data provide additional information for the breeder to consider in selecting a greenbug-resistant breeding line.     

Genetic mapping revealed two loci for soybean aphid resistance in PI 567301B

The soybean aphid (Aphis glycines Matsumura) is the most damaging insect pest of soybean [Glycine max (L.) Merr.] in North America. New soybean aphid biotypes have been evolving quickly and at least three confirmed biotypes have been reported in USA. These biotypes are capable of defeating most known aphid resistant soybean genes indicating the need for identification of new genes. Plant Introduction (PI) 567301B was earlier identified to have antixenosis resistance against biotype 1 and 2 of the soybean aphid. Two hundred and three F_7:9 recombinant inbred lines (RILs) developed from a cross of soybean aphid susceptible cultivar Wyandot and resistant PI 567301B were used for mapping aphid resistance genes using the quantitative trait loci (QTL) mapping approach. A subset of 94 RILs and 516 polymorphic SNP makers were used to construct a genome-wide molecular linkage map. Two candidate QTL regions for aphid resistance were identified on this linkage map. Fine mapping of the QTL regions was conducted with SSR markers using all 203 RILs. A major gene on chromosome 13 was mapped near the previously identified Rag2 gene. However, an earlier study revealed that the detached leaves of PI 567301B had no resistance against the soybean aphids while the detached leaves of PI 243540 (source of Rag2) maintained aphid resistance. These results and the earlier finding that PI 243540 showed antibiosis resistance and PI 567301B showed antixenosis type resistance, indicating that the aphid resistances in the two PIs are not controlled by the same gene. Thus, we have mapped a new gene near the Rag2 locus for soybean aphid resistance that should be useful in breeding for new aphid-resistant soybean cultivars. Molecular markers closely linked to this gene are available for marker-assisted breeding. Also, the minor locus found on chromosome 8 represents the first reported soybean aphid-resistant locus on this chromosome.     

Effect of cabbage cultivars with varying levels of resistance to aphids on the performance of the parasitoid,Aphidius colemani (Hymenoptera: Braconidae)

The effect of aphid resistance in plant cultivars on parasitism of Myzus persicae (Sulzer) and Brevicoryne brassicae (Linnaeus) by Aphidius colemani Viereck was investigated under laboratory conditions using three cultivars of common cabbage, Brassica oleracea var. capitata. Significantly greater aphid populations were found on cv. Derby Day (green-leaved, susceptible to both aphid species), regardless of the presence or absence of parasitoids, compared with cv. Minicole (green-leaved, partially resistant with antibiosis factors for B. brassicae) or cv. Ruby Ball (red-leaved, partially resistant with antixenosis factors for B. brassicae). Minicole had the greatest proportion of parasitized aphids and Derby Day the least. A significantly lower percentage of emerged parasitoids were recorded on Minicole for both aphid species compared with Derby Day and Ruby Ball. The implication of the results are discussed in relation to the integrated control of aphids.     

Antibiosis and tolerance to five species of spittlebug (Homoptera: Cercopidae) in Brachiaria spp.: implications for breeding for resistance

Several genera and species of spittlebugs (Homoptera: Cercopidae) are economic pests of Brachiaria spp. grasses in tropical America. To support current breeding programs aimed at obtaining multiple spittlebug resistance, we undertook a series of studies on antibiosis and tolerance as possible mechanisms of resistance to five major spittlebug species affecting Brachiaria spp. in Colombia: Aeneolamia varia (F.), Aeneolamia reducta (Lallemand), Zulia carbonaria (Lallemand), Zulia pubescens (F.), and Mahanarva trifissa (Jacobi). Four host genotypes, well known for their reaction to A. varia attack, were used to compare their resistance to other spittlebug species: CIAT 0654 and CIAT 0606 (susceptible) and CIAT 6294 and CIAT 36062 (resistant). CIAT 0654 and CIAT 36062 were used in antibiosis studies. Tolerance studies were conducted with CIAT 0654, CIAT 6294, and CIAT 36062. Sixty-five hybrid-derived clones were used to identify levels of multiple resistance to three spittlebug species. The levels of antibiosis resistance in CIAT 36062 clearly differed by spittlebug species and were classified as follows: very high for M. trifissa, high for A. varia and A. reducta, moderate for Z. pubescens, and absent for Z. carbonaria. Our results suggest the presence of true tolerance to Z. carbonaria in CIAT 6294 and CIAT 36062, true tolerance to Z. pubescens in CIAT 6294 and a combination of tolerance and antibiosis as mechanisms of resistance to Z. pubescens in CIAT 36062. Of the 65 hybrid clones tested with A. varia, A. reducta, and Z. carbonaria, 15 combined resistance to two species and three showed antibiosis resistance to all three spittlebug species.     

An investigation of the resistance to cabbage aphid in brassica species

Fifteen Brassica species and subspecies, as well as accessions of Arabidopsis thaliana and Eruca sativa, were evaluated for their resistance to the cabbage aphid, Brevicoryne brassicae, in the field and laboratory at Horticulture Research International (HRI) Wellesbourne in 1992. In the laboratory, aphid performance was measured in terms of pre-reproductive period, reproductive period, population increase and insect survival. Using these parameters it was possible to identify brassicas possessing antibiosis resistance. In the field B. brassicae attack was assessed in terms of numbers of insects and it was possible to determine aphid preferences for alighting on different species. High levels of antixenosis and antibiosis resistance were discovered in accessions of Brassica fruticulosa and B. spinescens, in a Brassica juncea breeding line and in Eruca sativa. Partial resistance was found in several other Brassica species. The potential of these various sources as donors of resistance genes to be bred into cultivated brassicas is discussed.     

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.     

Differences in susceptibility to attack by Brevicoryne brassicae (L.) on brussels sprouts

The attack by alate and other forms of Brevicoryne brassicae (L.) and the amount of leaf malformation they caused on each of eighteen varieties or plant breeders' lines of brussels sprout were assessed in both 1967 and 1968 over the first 6 weeks following planting out. The results of the two years agreed closely despite differences in the dates of planting relative to aphid immigration. As a group, the Dutch varieties Roodnerf, Gravendeel, Tribune, Hercules and Castricum Glory were preferred by the incoming alates but suffered less damage than the least preferred inbred line, 3BL, of Cambridge No. 3. Some evidence of antibiosis was obtained, especially from a later experiment with three Cambridge inbred lines. However, in 1969 when there was a high rate of aphid multiplication on six varieties representing sprout types from Holland, Cambridge, Bedford and Ireland, any expression of host non-preference or antibiosis was soon obscured by the development of large aphid infestations and much damage. Under these circumstances, considerable varietal differences in tolerance to attack were found. Irish Elegance sustained most damage yet its most heavily attacked plants supported little more than half of the aphid numbers of equivalent plants of the most tolerant variety Darkmar 21. The Dutch varieties Vremo Inter and Seven Hills were almost as tolerant as Darkmar 21 and recovered quickly from damage. Darkmar 21 also showed high tolerance to attack in 1970, as did the Dutch varieties Rollo, Roodnerf and Stiekema, when large aphid infestations developed on the eight varieties then grown. Lack of plant tolerance to damage appeared to be associated with incurling foliage, a characteristic of Irish Elegance, Cambridge No. 1 and No. 3.     

Characterization of greenbug (Homoptera: Aphididae) resistance in synthetic hexaploid wheats

Twelve greenbug (Schizaphis graminum (Rondani)) biotype E-resistant synthetic hexaploid wheats synthesized by crossing Triticum dicoccum Schrank. and Aegilops tauschii (Coss.) Schmal. were evaluated for the three known insect resistance categories, including antibiosis, anti-xenosis, and tolerance. Different methods were evaluated for calculating antibiosis and tolerance. Calculating intrinsic rate of population increase and measuring leaf chlorophyll content with a SPAD chlorophyll meter proved to be time- and labor-efficient for antibiosis and tolerance determination, respectively. The resistance in all synthetic hexaploids proved to be the result of a combination of antibiosis, antixenosis, and tolerance, which makes them valuable sources of greenbug resistance. To assist plant breeders in selecting the best germplasm for greenbug resistance, a plant resistance index was created that revealed differences among the synthetic hexaploid wheats.     

Co-localisation of host plant resistance QTLs affecting the performance and feeding behaviour of the aphid Myzus persicae in the peach tree

The architecture and action of quantitative trait loci (QTL) contributing to plant resistance mechanisms against aphids, the largest group of phloem-feeding insects, are not well understood. Comparative mapping of several components of resistance to the green peach aphid (Myzus persicae) was undertaken in Prunus davidiana, a wild species related to peach. An interspecific F(1) population of Prunus persica var. Summergrand × P. davidiana clone P1908 was scored for resistance (aphid colony development and foliar damage) and 17 aphid feeding behaviour traits monitored by means of the electrical penetration graph technique. Seven resistance QTLs were detected, individually explaining 6.1-43.1% of the phenotypic variation. Consistency was shown over several trials. Nine QTLs affecting aphid feeding behaviour were identified. All resistance QTLs except one co-located with QTLs underlying aphid feeding behaviour. A P. davidiana resistance allele at the major QTL was associated with drastic reductions in phloem sap ingestion by aphids, suggesting a phloem-based resistance mechanism. Resistance was also positively correlated with aphid salivation into sieve elements, suggesting an insect response to restore the appropriate conditions for ingestion after phloem occlusion. No significant QTL was found for traits characterising aphid mouthpart activity in plant tissues other than phloem vessels. Two QTLs with effects on aphid feeding behaviour but without effect on resistance were identified. SSR markers linked to the main QTLs involved in resistance are of potential use in marker-assisted selection for aphid resistance. Linking our results with the recent sequencing of the peach genome may help clarify the physiological resistance mechanisms.     

QTL analysis for aphid resistance and growth traits in apple

The rosy apple aphid (Dysaphis plantaginea), the leaf-curling aphid (Dysaphis cf. devecta) and the green apple aphid (Aphis pomi) are widespread pest insects that reduce growth of leaves, fruits and shoots in apple (Malus × domestica). Aphid control in apple orchards is generally achieved by insecticides, but alternative management options like growing resistant cultivars are needed for a more sustainable integrated pest management (IPM). A linkage map available for a segregating F₁-cross of the apple cultivars ‘Fiesta’ and ‘Discovery’ was used to investigate the genetic basis of resistance to aphids. Aphid infestation and plant growth characteristics were repeatedly assessed for the same 160 apple genotypes in three different environments and 2 consecutive years. We identified amplified fragment length polymorphism (AFLP) markers linked to quantitative trait loci (QTLs) for resistance to D. plantaginea (‘Fiesta’ linkage group 17, locus 57.7, marker E33M35–0269; heritability: 28.3%), and to D. cf. devecta (‘Fiesta’ linkage group 7, locus 4.5, marker E32M39–0195; heritability: 50.2%). Interactions between aphid species, differences in climatic conditions and the spatial distribution of aphid infestation were identified as possible factors impeding the detection of QTLs. A pedigree analysis of simple sequence repeat (SSR) marker alleles closely associated with the QTL markers revealed the presence of the alleles in other apple cultivars with reported aphid resistance (‘Wagener’, ‘Cox’s Orange Pippin’), highlighting the genetic basis and also the potential for gene pyramiding of aphid resistance in apple. Finally, significant QTLs for shoot length and stem diameter were identified, while there was no relationship between aphid resistance and plant trait QTLs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sources of variation in the interaction between three cereal aphids (Hemiptera: Aphididae) and wheat (Poaceae)

The relative contributions of host plant, herbivore species and clone to variation in the interaction between cereal aphids and wheat were investigated using five clones each of three species, Rhopalosiphum padi (Linnaeus), Sitobion avenae (Fabricius) and Schizaphis graminum (Rondani), on seedlings of two cultivars of Triticum aestivum L. and one cultivar of Triticum durum Desf. More individuals and biomass of R. padi than of the other two species were produced on seedlings. The three wheat cultivars lost similar amounts of biomass as a result of infestation by aphids, with the amount lost depending on aphid species: S. avenae caused the lowest loss in biomass. Variation in aphid biomass production was due mostly to differences among aphid species (70%), less to the interaction between wheat type and aphid species (7%), and least to aphid clone (1%). The specific impact of the aphids on the plants ranged from 1.7 to 3.7 mg of plant biomass lost per mg of aphid biomass gained, being lowest for R. padi and highest for S. graminum. Variation in plant biomass lost to herbivory was due mostly to unknown sources of error (95%), probably phenotypic differences among individual seedlings, with 3% due to aphid species and none attributable to aphid clone. For these aphid-plant interactions, differences among aphid clones within species contributed little to variation in aphid and plant productivity; therefore, a small sample of clones was adequate for quantifying the interactions. Furthermore, one clone of each species maintained in the laboratory for about 200 parthenogenetic generations was indistinguishable from clones collected recently from the field.