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.     

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.     

Induced resistance by Myzus persicae in the peach cultivar `Rubira'

The effect of a previous infestation by the green peach aphid Myzus persicae (Sulzer) on the settling behaviour and reproduction of the same aphid species was investigated in the resistant peach cultivar Rubira, and compared with that observed in the susceptible control cultivar GF305. A previous infestation of 48 h triggered induced resistance in Rubira. There were significantly fewer aphids settling on preinfested than on uninfested plants, indicating an increased rejection of Rubira as a host plant. The level of induced resistance in preinfested plants was positively related to the duration of the first infestation. In GF305, previous infestation had no detrimental effect on aphid settlement and even slightly enhanced larviposition by adult females. The aphid probing behaviour after a 48-h preinfestation was also monitored for 8 h with the electrical peneration graph (EPG) technique. On preinfested GF305, most EPG parameters indicated an enhanced host plant acceptance. On preinfested GF305, aphids produced less sieve element salivation and more continuous sap ingestion than on uninfested GF305, indicating that the previous aphids provoked changes in plant properties beneficial to the test aphids. In Rubira, a major induced factor of resistance was thought to be expressed in the sieve element as phloem sap ingestion was 4-fold shorter on preinfested than on uninfested plants. The time taken by the aphid stylets to reach a sieve element was also significantly increased on preinfested Rubira, suggesting the induction of resistance factors outside the phloem. The originality of the Rubira/M. persicae interaction is discussed in the perspective of a better understanding of plant induced responses to aphids.     

The potential of Prunus davidiana for introgression into peach [Prunus persica (L.) Batsch] assessed by comparative mapping

The potential for introgression of Prunus davidiana, a wild species related to peach, was evaluated with respect to problems of non-Mendelian segregation or suppressed recombination which often hamper breeding processes based on interspecific crosses. Three connected (F1, F2 and BC2) populations, derived from a cross between P. davidiana clone P1908 and the peach cultivar Summergrand were used. The intraspecific map of P. davidiana already established using the F1 progeny was complemented, and two interspecific maps, for the F2 and BC2 progenies, were built with a set of markers selected from the Prunus reference map. With the molecular data collected for the F2 map construction, regions with distorted marker segregation were detected on the genome; one third of all loci deviated significantly from the expected Mendelian ratios. However, some of these distorted segregations were probably not due to the interspecific cross. On linkage group 6, a skewed area under gametic selection was most likely influenced by the self-incompatibility gene of P. davidiana. Using anchor loci, a good colinearity between the three maps built and the Prunus reference map was demonstrated. Comparative mapping also revealed that homologous recombination occurred normally between P. davidiana and the Prunus persica genome. This confirmed the closeness of the two species. Higher recombination rates were generally observed between P. davidiana and P. persica than between Prunus amygdalus and P. persica. The consequences for plant breeding strategy are discussed. The three maps of the F1, F2 and BC2 progenies provide useful tools for QTL detection and marker-assisted selection, as well as for assessing the efficiency of the peach breeding scheme applied to introgress P. davidiana genes into peach cultivated varieties.     

Quantitative trait loci analysis of <Emphasis Type="Italic">Plum pox virus</Emphasis> resistance in <Emphasis Type="Italic">Prunus davidiana</Emphasis> P1908: new insights on the organization of genomic resistance regions

No valuable source of resistance to Plum pox virus (PPV), the causative agent of sharka disease, has been found in peach (Prunus persica), but polygenic resistance to PPV was described in Prunus davidiana clone P1908. Two previous studies using F₁ and F₂ populations derived from the nectarine cv. Summergrand and P. davidiana P1908 identified a total of six P. davidiana quantitative trait loci (QTLs) involved in PPV resistance (Marcus strain). In an effort to verify the QTL stability in a large progeny and to search for possible interactions of the genetic backgrounds, the current study evaluated the incidence of PPV infection in an F₁ population derived from the susceptible peach cv. Rubira and P. davidiana P1908 over three growth periods using an improved method of PPV phenotyping referred to as “heavy test.” The phenotypic dataset was analyzed using similar methods as the previous studies and a newly developed simple-sequence-repeat-based P. davidiana map. Nine regions involved in differential symptom expression were identified among which six were common between studies. However, the level of resistance observed in the population was very low compared to the other studies, and the main QTL previously identified in linkage group 6 was not conserved, suggesting strong interaction of the genetic background of the susceptible parent with that of P. davidiana 1908. Consequently, this could be a limiting factor for developing resistant cultivars derived from P. davidiana P1908.     

Differences in Strain Virulence of the European Stone Fruit Yellows Phytoplasma and Susceptibility of Stone Fruit Trees on Various Rootstocks to this Pathogen

Twenty strains of the European stone fruit yellows (ESFY) phytoplasma showed great differences in virulence when examined by graft inoculation of trees on peach, peach hybrid GF 677 and P. 'Marianna' GF 8/1 rootstocks. The most virulent strains killed all trees on peach rootstocks whereas the mild strains did not cause mortality but induced only mild foliar symptoms and slightly reduced vigour. Virulence often depended on the pathogen–scion combination and was in several cases most severe when the scion consisted of the original host of the pathogen. To examine resistance in stone fruits, trees on a total of 23 rootstocks were inoculated with the ESFY strains. Trees on the Prunus domestica stocks Ackermann's, Brompton and P 1275 and on Prunus cerasifera stock Myrabi were little affected. Slightly more damage occurred in trees on rootstocks GF 677, GF 8–1, and the P. insititia stocks St Julien A and St Julien GF 655/2. Ishtara, P. cerasifera stock Myrobalan, and peach rootstocks Higama and GF 305 were shown to be moderately susceptible and a high susceptibility was found in trees on peach rootstocks Montclar, peach seedling, Rutgers Red Leaf, and Rubira, on apricot seedlings and St Julien 2. Of flowering cherry trees on various rootstocks, the least susceptible were those on Gisela 3 and F 12/1 whereas Gisela 1, Weihroot 158 and Gisela 5 were more affected. Phytoplasmas were detected by either DAPI (4'-6-diamidino-2-phenylindole) staining or polymerase chain reaction in all rootstocks and scions tested. However, detection frequency and phytoplasma concentrations were usually lower in the more tolerant hosts than in susceptible genotypes.     

S-adenosylmethionine synthase (BtSAMS) from Balsas teosinte confers resistance to peach aphids in Arabidopsis

The purpose of this study was to clone S-adenosylmethionine synthase (BtSAMS) from Balsas teosinte (Zea mays ssp. parviglumis) and investigate its function via heterologous expression in Arabidopsis plants. The BtSAMS coding sequence was cloned and its length is 1185 bp. The expression of BtSAMS in Z. mays ssp. parviglumis was a 12.2-fold increase after aphid infestation. Transgenic Arabidopsis plants heterologously expressing BtSAMS were generated and their morphological characters were observed. A prolonged vegetative growth period was displayed by BtSAMS-OX plants. The more and longer trichomes were observed on the upper surface of BtSAMS seedlings. The ethylene content in BtSAMS-OX plants was not significantly different compared with the untransformed controls. These BtSAMS-OX transgenic plants showed greater resistance to peach aphids compared with the untransformed control plants. Peach aphids appeared to prefer the untransformed controls in that aphids number reduced by more than 50% on BtSAMS plants compared to controls. When sprayed with the insecticide avi-imidacloprid, the number of peach aphids decreased on BtSAMS-OX plants and untransformed controls by 91.6% and 80.5%, respectively. Peach aphids also showed lower reproduction ability when forced to feed on the transgenic plants. The number of newly born nymphs on BtSAMS plants was less than 50% of that on the untransformed control. We therefore propose that BtSAMS may be a suitable candidate gene to confer resistance to peach aphids in plants.     

桃蚜取食行为对大气CO_2浓度升高的响应

【目的】近年来随着人类活动的增加,温室气体尤其是大气CO_2浓度升高造成的虫害爆发已成为国际上关注的焦点,因此,研究拟南芥Arabidopsis thaliana上桃蚜取食行为的变化对大气CO_2浓度升高的响应意义重大。【方法】本研究以拟南芥和绿色桃蚜Myzus persicae为研究对象,利用野生型拟南芥Col-0,茉莉酸途径信号传导缺失突变体(jar1)、水杨酸途径信号传导缺失突变体(npr1)、乙烯途径信号传导缺失突变体(ein2-5)为材料,以大气CO_2浓度升高为影响因子,利用刺吸式电位仪(EPG)记录了桃蚜在不同处理的拟南芥上的取食波形。【结果】研究结果发现:CO_2浓度升高缩短了Col-0和jar1植株上蚜虫首次刺探时间和首次到达韧皮部的时间,却延长了npr1和ein2-5上蚜虫首次到达韧皮部的时间,降低了jar1植株上蚜虫总的刺探时间且增加了其总的取食韧皮部时间,但没有改变其它基因型植株上蚜虫总的刺探和取食时间;同时增加了野生型植株上蚜虫的刺探频率,却没有影响其它基因型植株上的刺探频率。【结论】CO_2浓度升高降低了野生型植株和jar1植株抗性,有利于蚜虫到达韧皮部;却增加了npr1和ein2-5上的植物抗性,从而不利于蚜虫到达韧皮部。     

Drought‐stress and plant resistance affect herbivore performance and proteome: the case of the green peach aphid Myzus persicae (Hemiptera: Aphididae)

Little is known about the simultaneous effects of drought stress and plant resistance on herbivorous insects. By subjecting the green peach aphid Myzus persicae Sulzer to well‐watered and drought‐stressed plants of both susceptible and resistant peach (Prunus persica), the effects of both stressors on aphid performance and proteomics are tested. Overall, the influence of the water treatment on aphid performance is less pronounced than the effect of host plant genetic resistance. On the susceptible cultivar, aphid survival, host acceptance and ability to colonize the plant do not depend on water treatment. On the resistant cultivar, aphid survival and ability to colonize are higher on drought‐stressed than on well‐watered plants. A study examining the pattern of protein expression aiming to explain the variation in aphid performance finds higher protein expression in aphids on the drought‐stressed susceptible cultivars compared with the well‐watered ones. In the susceptible cultivar, the regulated proteins are related to energy metabolism and exoskeleton functionality, whereas, in the resistant cultivar, the proteins are involved with the cytoskeleton. Comparison of the protein expression ratios for resistant versus susceptible plants reveals that four proteins are down‐regulated in well‐watered plants and 15 proteins are down‐regulated in drought‐stressed plants. Drought stress applied to the susceptible cultivar induces the regulation of proteins in M. persicae that enable physiological adaptation to maintain an almost unaltered aphid performance. By contrast, for aphids on the resistant cultivar subjected to drought stress, the down‐regulation of proteins responds to an induced host susceptibility effect.     

Mapping Rm2 gene conferring resistance to the green peach aphid (Myzus persicae Sulzer) in the peach cultivar ��Rubira?��

The green peach aphid (GPA), Myzus persicae (Sulzer), is a widespread pest insect that significantly reduces yield in peach orchards [Prunus persica (L.) Batsch]. Chemical control of the GPA population in the orchards showed little efficiency because of the development of resistance to most classes of insecticides. Biological control partially gave convincing results. Breeding for resistant peach cultivars is therefore a serious option to take into account for the development of sustainable pest management. Among the few available resistance cultivars, the rootstock peach “Rubira?” shows a strong induced antixenosis-type GPA resistance. This was demonstrated segregating as a single dominant gene. In order to investigate the genetic basis of resistance and develop molecular tools useful in breeding programs, a F₂ population derived from “Rubira?” also segregating for leaf color was grown and scored for GPA resistance under contrasted environmental conditions. An SSR-based genetic linkage map composed of 120 SSR loci spanned over a distance of 497.8 cM was then established. The GPA resistance mapped to a single locus at the bottom end of linkage group 1. We propose to name Rm2 the dominant allele of the underlying gene. Additionally, a reciprocal translocation was identified near the Gr gene controlling leaf color. The red-leaf parent “Rubira?” was demonstrated responsible for the translocation. This study provides the basis for future molecular analysis for the use of Rm2 in peach breeding programs against GPA in peach orchards.     

Antibiosis and antixenosis of six commonly produced potato cultivars to the green peach aphid, Myzus persicae Sulzer (Hemiptera: Aphididae)

The antibiotic and antixenotic resistance of six commonly produced potato cultivars in Iran including Aozonia, Agria, Cosima, Cosmos, Kondor and Savalan to the green peach aphid, Myzus persicae Sulzer, were investigated under laboratory conditions at 20 ± 2oC, 65 ± 5% RH and 16:8h (L:D) in 2009. Antibiosis experiments showed significant differences in the developmental time, nymphal survivorship, fecundity, adult longevity of the green peach aphid among the potato cultivars. Intrinsic rate of natural increase (r m) for apterous aphids varied significantly with the potato cultivars on which the aphids were reared. This value ranged from 0.225 to 0.293 females/female/day, which was lowest on Cosmos and highest on Aozonia. Additionally, the estimated net reproductive rate (R0) and finite rate of increase (λ) for apterous aphids were the lowest on Cosmos. For the antixenosis experiment, no significant difference was found in aphid's preference to the potato cultivars. However, Aozonia was preferred more than the other five cultivars by the apterous aphids. Therefore, our results demonstrated that among the investigated cultivars the Cosmos cultivar is moderately resistant to the green peach aphid.     

QTLs for powdery mildew resistance in peach ×Prunus davidiana crosses: consistency across generations and environments

Powdery mildew, caused by Sphaerotheca pannosa var. persicae is one of the most important diseases in European peach orchards. Quantitative trait loci controlling powdery mildew resistance were detected using three related F1, F2 and BC2 populations derived from the cross between the resistant parent P. davidiana clone P1908 and the susceptible peach cultivar Summergrand. Powdery mildew resistance of each population was evaluated under natural exposure, in several locations and over several years. Thirteen QTLs were detected. For nine of them, the favourable allele came from the resistant parent. Five QTLs were consistently detected across the three populations. The F1 hybrid used to produce F2 and BC2 populations had not inherited the favourable allele from P1908 for QTL detected on LG3 and LG8 in F1 population. QTLs were not detected in the corresponding regions in F2 and BC2 populations. In two other genomic areas, significant substitution effects between P1908 alleles were evidenced in the F1 population, but the favourable allele came from Summergrand in the F2 and BC2 populations. Analysis of phenotypic data suggested an important qualitative change in the distribution of powdery mildew resistance after 1996, confirmed by QTL analysis. Indeed, a dramatic decrease of the effect of the major QTL previously detected on LG6 was observed after 1996, while the QTL on LG8 was increasingly involved in the control of powdery mildew resistance. Consequences for peach breeding strategies to improve powdery mildew resistance are discussed.     

3种寄主上桃蚜的选择性及形态分化

桃蚜是一种重要的农业害虫,寄主广泛,种下分化复杂。以采自黄土高原旱作区桃树、烟草、甘蓝上的桃蚜为研究对象,通过叶片选择法、传统比较形态测定法研究了3种寄主上桃蚜的选择性及形态分化。结果表明:在3种寄主同时存在的情况下,烟草上的桃蚜嗜食烟草,表现为63.5%的桃蚜选择烟草叶,13.8%选择甘蓝叶,8.2%选择桃叶,而甘蓝和桃树上的桃蚜对原寄主没有表现出明显的嗜好性;从形态指标来看,3种寄主上的桃蚜在体长、触角末节长度、后足腿节长度、触角与体长的比例方面存在显著差异(P0.05),说明这几个特征可以作为区分这3种寄主上桃蚜的依据。综合分析可以初步认为黄土高原旱作区烟草上的桃蚜可能形成了寄主专化型-烟草型。     

Resistance induction and herbivore virulence in the interaction between Myzus persicae (Sulzer) and a major aphid resistance gene (Rm2) from peach

In gene-for-gene host–enemy interactions, monogenic plant resistance results from pathogen recognition that initiates the induction of plant defense responses. Schematically, as the result of the on/off process of recognition, phenotypic variability in enemy virulence is expected to be qualitative, with either a failure or a success of host colonization. We focussed on a major gene from peach conferring avoidance resistance against the green peach aphid Myzus persicae. Measurements of herbivore density and time-dependent aspects of resistance induction were examined, as well as variability in the aphid’s ability to exploit the resistant host. Varying densities of infestation did not provoke differences in the aphid’s tendency to leave a plant, and a single aphid was sufficient to elicit a response. Similarly, the duration of infestation did not affect the aphid response. A brief aphid feeding time of 3 h triggered induced resistance, which became effective between 24 and 48 h after the initial attack. Induced resistance decayed over time in the absence of additional infestation. Thirty aphid genotypes collected from natural populations were tested in the laboratory. No clone could colonize the resistant host, suggesting that all of them triggered the induction of effective plant defense responses. However, we detected significant quantitative variation among clones in the tendency of aphids to leave plants. These results improve our understanding of induced resistance as a dynamic phenomenon and suggest that the potential for aphids to adapt to a major plant resistance gene may depend on factors other than the mere capacity to evade recognition.     

Analogues of virus resistance genes map to QTLs for resistance to sharka disease in <Emphasis Type="Italic">Prunus davidiana</Emphasis>

Plum pox virus (PPV), the causative agent of sharka disease in Prunoideae, is one of the most serious problems affecting stone fruit production in Europe and America. Resistance to PPV was previously described in a Prunus davidiana clone, P1908, and introduced into peach (Prunus persica) genotypes. Genetic resistance to PPV displays a complex pattern of quantitative inheritance. An analysis of quantitative trait loci (QTLs) for resistance was performed on an F1 interspecific peach population obtained from a cross between the susceptible nectarine cultivar Summergrand and P. davidiana. The hybrids were graft-inoculated with PPV in duplicate following a classical procedure. The incidence of infection was evaluated four times, over two vegetative cycles, by symptom observation and enzyme-linked immunoadsorbent assays (ELISA). Restriction of systemic downward movement of the PPV virus was also evaluated by testing the susceptible rootstocks. Using both analysis of variance and non-parametric tests, six genomic regions involved in PPV resistance were detected. Depending on the scoring data considered, between 22 and 51% of the phenotypic variance could be explained by the quantitative model. One QTL, located in the distal region of linkage group 1, maps in a genomic region that is syntenic to the location of a resistance gene previously identified in the apricot cv. Goldrich. Some QTLs appeared to be temporally specific, reflecting the environmental dependence of PPV-resistance scoring. Candidate gene fragments were amplified by PCR, isolated and mapped on the peach interspecific linkage map. We report here the co-localization of three analogues of virus resistance genes with two distinct genomic regions linked to PPV resistance in P. davidiana.Electronic Supplementary Material Supplementary material is available for this article at     

Nitrogen and water supplies affect peach tree–green peach aphid interactions: the key role played by vegetative growth

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烟蚜γ_m和排蜜量之间的关系(英文)

1990—1992年进行了烟蚜在50个烟草品种上取食的r_m和排蜜量与田间蚜量之间关系的研究。结果表明:r_m的变化范围为0.102～0.498,烟蚜24小时的排蜜量变化范围为3.862～29.356mm~2/头,并由此说明烟草品种对烟蚜的抗性是由于降低了其净生殖率。r_m和排蜜量与田间蚜量有显著相关关系,因而可用排蜜量来作为烟草品种抗蚜性的一个鉴定指标。     

Control of Myzus persicae and Lipaphis erysimi (Hemiptera: Aphididae) by adults and larvae of a flightless strain of Harmonia axyridis (Coleoptera: Coccinellidae) on non-heading Brassica cultivars in the greenhouse

The green peach aphid [Myzus persicae (Sulzer)] and turnip aphid [Lipaphis erysimi (Kaltenbach)] are economically important pests with a worldwide distribution. We have evaluated the efficacy of releasing adults and larvae of a flightless strain of the multicolored Asian lady beetle (Harmonia axyridis Pallas) as a control measure against these aphids on plants of non-heading Brassica cultivars. Both adults and larvae of H. axyridis were observed to be effective biocontrol agents, markedly decreasing the numbers of aphids. The residence duration of adults was longer than that of larvae. The proportion of non-marketable plants damaged by aphids was lower in plots into which either adults or adults and larvae of H. axyridis had been released. These results suggest that both adults and larvae of this flightless strain of H. axyridis are effective in controlling aphids on plants of non-heading Brassica cultivars.     

人工饲料pH值对桃蚜存活、繁殖及抗氧化酶活性的影响

通过测定在不同pH值饲料上桃蚜Myzus persicae（Sulzer）的存活率、繁殖率及其抗氧化酶活性的变化,评价饲料pH值对桃蚜生态和生理指标的影响,筛选扩繁桃蚜饲料最适pH值。结果显示：不同pH值人工饲料饲喂桃蚜,其存活率3d后即呈现显著差异,在试验调查时间内,饲料pH值为7.0处理的存活率、单雌产蚜量和内禀增长率均为最高;不同处理桃蚜体内的超氧化物歧化酶（SOD）、过氧化氢酶（CAT）活性与对照处理相比均存在显著性差异,在24h左右达到最大值,而后有所下降;饲料pH值为7.0和7.2两处理的酶活性变化幅度较小,在48h后接近正常水平。试验结果表明,扩繁桃蚜较适合的人工饲料pH值为7.0左右。