Phloem-based resistance to green peach aphid is controlled by Arabidopsis PHYTOALEXIN DEFICIENT4 without its signaling partner ENHANCED DISEASE SUSCEPTIBILITY1

Green peach aphid (GPA) Myzus persicae (Sülzer) is a phloem-feeding insect with an exceptionally wide host range. Previously, it has been shown that Arabidopsis thaliana PHYTOALEXIN DEFICIENT4 (PAD4), which is expressed at elevated levels in response to GPA infestation, is required for resistance to GPA in the Arabidopsis accession Columbia. We demonstrate here that the role of PAD4 in the response to GPA is conserved in Arabidopsis accessions Wassilewskija and Landsberg erecta. Electrical monitoring of aphid feeding behavior revealed that PAD4 modulates a phloem-based defense mechanism against GPA. GPA spends more time actively feeding from the sieve elements of pad4 mutants than from wild-type plants, and less time feeding on transgenic plants in which PAD4 is ectopically expressed. The activity of PAD4 in limiting phloem sap uptake serves as a deterrent in host-plant choice, and restricts aphid population size. In Arabidopsis defense against pathogens, all known PAD4 functions require its signaling and stabilizing partner EDS1 (ENHANCED DISEASE SUSCEPTIBILITY1). Bioassays with eds1 mutants alone or in combination with pad4 and with plants conditionally expressing PAD4 under the control of a dexamethasone-inducible promoter reveal that PAD4-modulated defense against GPA does not involve EDS1. Thus, a PAD4 mode of action that is uncoupled from EDS1 determines the extent of aphid feeding in the phloem.     

Redox responses of Arabidopsis thaliana to the green peach aphid,Myzus persicae

The green peach aphid (Myzus persicae) is a phloem-feeding insect that causes economic damage on a wide array of crops. Using a luminol-based assay, a superoxide-responsive reporter gene (Zat12::luciferase), and a probe specific to hydrogen peroxide (HyPer), we demonstrated that this aphid induces accumulation of reactive oxygen species (ROS) in Arabidopsis thaliana. Similar to the apoplastic oxidative burst induced by pathogens, this response to aphids was rapid and transient, with two peaks occurring within 1 and 4 hr after infestation. Aphid infestation also induced an oxidative response in the cytosol and peroxisomes, as measured using a redox-sensitive variant of green fluorescent protein (roGFP2). This intracellular response began within minutes of infestation but persisted 20 hr or more after inoculation, and the response of the peroxisomes appeared stronger than the response in the cytosol. Our results suggest that the oxidative response to aphids involves both apoplastic and intracellular sources of ROS, including ROS generation in the peroxisomes, and these different sources of ROS may potentially differ in their impacts on host suitability for aphids.     

Myzus persicae (green peach aphid) feeding on Arabidopsis induces the formation of a deterrent indole glucosinolate

Cruciferous plants produce a wide variety of glucosinolates as a protection against herbivores and pathogens. However, very little is known about the importance of individual glucosinolates in plant defense and the regulation of their production in response to herbivory. When Myzus persicae (green peach aphid) feeds on Arabidopsis aliphatic glucosinolates pass through the aphid gut intact, but indole glucosinolates are mostly degraded. Although aphid feeding causes an overall decrease in Arabidopsis glucosinolate content, the production of 4-methoxyindol-3-ylmethylglucosinolate is induced. This altered glucosinolate profile is not a systemic plant response, but is limited to the area in which aphids are feeding. Aphid feeding on detached leaves causes a similar change in the glucosinolate profile, demonstrating that glucosinolate transport is not required for the observed changes. Salicylate-mediated signaling has been implicated in other plant responses to aphid feeding. However, analysis of eds5, pad4, npr1 and NahG transgenic Arabidopsis, which are compromised in this pathway, demonstrated that aphid-induced changes in the indole glucosinolate profile were unaffected. The addition of purified indol-3-ylmethylglucosinolate to the petioles of cyp79B2 cyp79B3 mutant leaves, which do not produce indole glucosinolates, showed that this glucosinolate serves as a precursor for the aphid-induced synthesis of 4-methoxyindol-3-ylmethylglucosinolate. In artificial diets, 4-methoxyindol-3-ylmethylglucosinolate is a significantly greater aphid deterrent in the absence of myrosinase than its metabolic precursor indol-3-ylmethylglucosinolate. Together, these results demonstrate that, in response to aphid feeding, Arabidopsis plants convert one indole glucosinolate to another that provides a greater defensive benefit.     

Tomato responds to green peach aphid infestation with the activation of trehalose metabolism and starch accumulation

The disaccharide trehalose and trehalose-6-phosphate that are present in trace amounts are suggested to have a signaling function in plants. Recently, it was demonstrated that trehalose metabolism contributes to Arabidopsis thaliana defense against the green peach aphid (GPA; Myzus persicae Sülzer), an important insect pest of a large variety of plants. TPS11 (TREHALOSE PHOSPHATE SYNTHASE11)-dependent trehalose metabolism was shown to curtail GPA infestation by promoting starch accumulation and expression of the PAD4 (PHYTOALEXIN-DEFICIENT4) gene, which has important roles in regulating antibiosis and antixenosis against GPA. Here we show that trehalose metabolism is similarly activated in leaves of GPA-infested tomato (Solanum lycopersicum) plants and likely contributes to tomato defense against GPA. GPA-infested leaves of tomato accumulated trehalose, which was accompanied by the transient upregulation of SlTPS11, a homolog of the Arabidopsis TPS11. GPA-infestation was also accompanied by starch accumulation and the upregulation of SlPAD4, the tomato homolog of Arabidopsis PAD4. Furthermore, trehalose application induced SlPAD4 expression and starch accumulation, and curtailed GPA infestation, suggesting that like in Arabidopsis trehalose contributes to tomato defense against GPA.     

桃蚜不同蚜型DNA多态性的RAPD研究

采用RAPD方法,对全周期桃蚜的有翅产雌性母蚜、无翅性母蚜、雄蚜、卵、干母、干雌、有翅迁移蚜等蚜型的DNA遗传多态性进行了分析。结果表明：卵的DNA多态性最大,性蚜次之,孤雌生殖蚜最小;卵与其它蚜型之间在遗传上具最大差异,其中与孤雌生蚜的差异大于与性蚜的;干母、干雌和迁移蚜之间的遗传关系最近。     

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.     

Studies on the morphometry of the green peach aphid, Myzus persicae (Sulzer) (Homoptera: Aphididae)

Aphids are one of the most important groups of phytophagous insects because of their polyphenism, host alternation, heteroecious behavior and reproductive habits. Laboratory culture of the aphid Myzus persicae on the host plant Brassica pekinensis Hubner was maintained seasonally for biometrical studies. The metric analysis revealed gradual developmental changes from the first instar nymph to the adult stage in size, shape, and a number of characters. The transitional changes were sufficiently distinct to allow differentiation of each of the life stages irrespective of the season. A key to the nymphal stages of the pest is provided.     

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

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Identification of indole glucosinolate breakdown products with antifeedant effects on Myzus persicae (green peach aphid)

The cleavage of glucosinolates by myrosinase to produce toxic breakdown products is a characteristic insect defense of cruciferous plants. Although green peach aphids ( Myzus persicae ) are able to avoid most contact with myrosinase when feeding from the phloem of Arabidopsis thaliana , indole glucosinolates are nevertheless degraded during passage through the insects. A defensive role for indole glucosinolates is suggested by the observation that atr1D mutant plants, which overproduce indole glucosinolates, are more resistant to M. persicae , whereas cyp79B2 cyp79B3 double mutants, which lack indole glucosinolates, succumb to M. persicae more rapidly. Indole glucosinolate breakdown products, including conjugates formed with ascorbate, glutathione and amino acids, are elevated in the honeydew of M. persicae feeding from atr1D mutant plants, but are absent when the aphids are feeding on cyp79B2 cyp79B3 double mutants. M. persicae feeding from wild-type plants and myrosinase-deficient tgg1 tgg2 double mutants excrete a similar profile of indole glucosinolate-derived metabolites, indicating that the breakdown is independent of these foliar myrosinases. Artificial diet experiments show that the reaction of indole-3-carbinol, a breakdown product of indol-3-ylmethylglucosinolate, with ascorbate, glutathione and cysteine produces diindolylmethylcysteines and other conjugates that have antifeedant effects on M. persicae . Therefore, the post-ingestive breakdown of indole glucosinolates provides a defense against herbivores such as aphids that can avoid glucosinolate activation by plant myrosinases.     

我国不同地区烟蚜种群生殖特征研究

研究了我国4个地区3种生活史类型的烟蚜在烟草上的种群繁殖特征,并用Weibull模型对种群存活特征和生殖特征进行了分析。结果表明:25±1℃、14L:10D、70%±10%RH条件下,长春地区烟蚜的生殖前期和产仔高峰日分别为大于等于7d和第11d,而南京、湖南郴州和河南宜阳烟蚜的生殖前期和产仔高峰日分别为小于7d和第9d。烟蚜种群的R0、T和rm在同一地区不同体色之间以及不同地区同一体色之间均存在有差异显著性,平均寿命仅在不同地区的红色烟蚜和褐色烟蚜中有显著性差异,在黄绿色之间差异均不显著。     

低浓度SO2污染与桃蚜的生长和繁殖

用40或80ppb SO_2熏气处理过的油菜饲养桃蚜(Myzus persicae),研究低浓度SO_2污染对这种蚜虫的影响。与对照组相比,两个处理组若蚜的发育速率显著加快,5天内的平均相对生长速率(MRGR)分别增加22.3％和31.5％。成蚜的繁殖力提高40—90％,导致桃蚜种群的内禀增长率(r_m)分别升高17％和27％。比较了桃蚜和其它3种蚜虫对SO_2污染的反应强度,讨论了SO_2污染促进某些植食性昆虫生长的可能原因。     

低浓度SO2污染与桃蚜的生长和繁殖

用40或80ppbSO₂熏气处理过的油菜饲养桃蚜(Myzus persicae),研究低浓度SO₂污染对这种蚜虫的影响。与对照组相比,两个处理组若蚜的发育速率显著加快,5天内的平均相对生长速率(MRGR)分别增加22.3%和31.5%。成蚜的繁殖力提高40—90%,导致桃蚜种群的内禀增长率(r_m)分别升高17%和27%。比较了桃蚜和其它3种蚜虫对SO₂污染的反应强度,讨论了SO₂污染促进某些植食性昆虫生长的可能原因。     

Settling behaviour and reproductive potential of the green peach aphid Myzus persicae on peach varieties and a related wild Prunus

Discovery in the late seventies of resistance to the green peach aphid Myzus persicae (Sulzer) (Homoptera: Aphididae) in Prunus species was based on screening in the field or in greenhouses with natural aphid populations. Here, we assess the impact of these wild and domesticated peach trees on the behaviour, development, reproductive performance and demography of cloned aphids under controlled light and temperature. Four peach varieties, i.e., Rubira, Weeping Flower Peach, Summergrand and Malo konare and the clone P1908 of the related species Prunus davidiana were tested against the highly susceptible cultivar GF305. Besides a variability in the performance of aphids among experiments, our results showed that (i) distinct mechanisms were involved in the sources of resistance studied and (ii) the ranking of the genotypes on their resistance/susceptibility status remained roughly stable throughout the experiments. Observations on the settling behaviour of first instar nymphs demonstrated antixenosis components in the resistance conferred by Rubira and Weeping Flower Peach. Nymphs began to leave the plants after a short exposure (19–21 h) and no aphid was left after 4 days. Nymphal mortality remained rather low (16%) compared to the repellent effect on aphids of both genotypes. Nymphs disappeared from Weeping Flower Peach significantly earlier than from Rubira. Summergrand, Malo konare and P. davidiana clone P1908 were accepted as host plants by aphids. On P. davidiana, decreased fecundity and intrinsic rate of natural increase (rm=0.20, averaged on all experiments) were clear expression of antibiosis. In addition, the mean length of the mature embryos within the gonads of the females on the day of adult moult was negatively correlated with the total number of embryos, providing evidence that aphids on this genotype lacked sufficient ressources to be directed both towards production and growth of embryos. Compared to the most susceptible cultivar GF305 (rm=0.36), Summergrand (rm=0.26) and Malo konare (rm=0.28) had, to a lesser extent, a negative impact on nymph production and rate of increase.     

Sublethal effects of selected insecticides on fecundity and wing dimorphism of green peach aphid (Hom., Aphididae)

Effects of sublethal concentrations (LC₂₅) of six insecticides (imidacloprid, rotenone, fenvalerate, abamectin, pirimicarb and azadirachtin) on fecundity and wing dimorphism of the green peach aphid, Myzus persicae (Sulzer), were studied both under laboratory and greenhouse conditions. In the laboratory, aphid reproduction reduced by 44.29% and 54.01% when rotenone and abamectin treatments were applied at sublethal dose, respectively, and sublethal fenvalerate application resulted in markedly lower average reproduction per female per day compared with control. Reproductive duration of aphid treated with abamectin significantly decreased by 44.19%. But in the greenhouse, there were no evident differences in the aphid fecundity and reproductive duration between treatments and control. Life‐table parameters also demonstrated that the six insecticides at sublethal doses did not stimulate the aphid reproductive potential. In the laboratory, after being exposed to sublethal doses of imidacloprid and fenvalerate, the proportions of alate progeny in aphid progeny were significantly higher than that of the control. In the greenhouse, percentages of alate offspring from the mother aphids treated with imidacloprid, fenvalerate and abamectin increased pronouncedly compared with control. Mortality rates of offspring in the nymphal stages from adults treated with insecticides revealed no significant changes between laboratory and greenhouse. The developmental time in days of the offspring varied in all treatments. Mechanisms of insecticide‐induced resurgence are discussed.     

桃蚜自然种群初级和次级共生菌的分子鉴定

蚜虫不仅带有专性初级内共生菌,还常常带有不同类群的兼性次级共生菌。本研究采用真细菌16S rDNA的通用引物,从桃蚜Myzus persicae (Sulzer)烟草种群体内扩增出1.5 kb的共迁移目的条带,然后将其克隆,并在阳性克隆筛选时进行了RFLP分析。当采用EcoRⅠ进行单酶切分析时,目的片段被切割成～650 bp和～850 bp两个小片段;当采用EcoRⅠ和HindⅢ进行双酶切分析时,该蚜群共生菌酶切谱带被分成2组,其中1组(GroupⅠ)只具有一个EcoRⅠ酶切位点,而另1组(Group Ⅱ)不仅具有一个EcoRⅠ酶切位点,还具有一个HindⅢ酶切位点,而且GroupⅠ为优势共生菌群。在此基础上分别对2组共生菌的16S rDNA全序列（～1.5 kb）进行了测定,结果表明：GroupⅠ属于泛菌属Pantoea,与成团泛菌Pantoea agglomerans亲缘关系最近（同源性达99.70%）,而Group Ⅱ属于蚜虫的专性内共生菌,即Buchnera aphidicola（同源性达99.50%）。这是在蚜虫体内存在泛菌次级共生菌的首次报道。     

Elevated CO2 shifts the focus of tobacco plant defences from cucumber mosaic virus to the green peach aphid

Elevation in CO₂ concentration broadly impacts plant physiological characteristics, which influences herbivores and biotrophic pathogens, which in turn regulate the plant defensive response. In this study, responses of tobacco plants to stress in the form of the green peach aphid, Myzus persicae (Sulzer), or cucumber mosaic virus (CMV), or both aphid and CMV combined were investigated in open‐top chambers under ambient and elevated CO₂ concentrations. We measured aboveground biomass and foliar chlorophyll, nitrogen, non‐structural carbohydrates, soluble protein, total amino acid and nicotine content in tobacco plants and also measured aphid population dynamics, body weight, honeydew production and anti‐oxidative enzyme activities in individual aphids. Plants produced more secondary metabolites for defence in both CO₂ treatments when treated with aphid and CMV combined than with either alone. Aphid density significantly increased on CMV‐infected tobacco plants (relative to uninfected plants) under ambient CO₂ but not under elevated CO₂. This suggests that plant defences against virus and aphid would be more efficient under elevated CO₂. Plant defence appears to shift from plant virus to aphid under increasing CO₂ levels, which highlights the potential influences of multiple biotic stressors on plants under elevated CO₂.     

TREHALOSE PHOSPHATE SYNTHASE11-dependent trehalose metabolism promotes Arabidopsis thaliana defense against the phloem-feeding insect Myzus persicae

Agricultural productivity is limited by the removal of sap, alterations in source-sink patterns, and viral diseases vectored by aphids, which are phloem-feeding pests. Here we show that TREHALOSE PHOSPHATE SYNTHASE11 (TPS11) gene-dependent trehalose metabolism regulates Arabidopsis thaliana defense against Myzus persicae (Sülzer), commonly known as the green peach aphid (GPA). GPA infestation of Arabidopsis resulted in a transient increase in trehalose and expression of the TPS11 gene, which encodes a trehalose-6-phosphate synthase/phosphatase. Knockout of TPS11 function abolished trehalose increases in GPA-infested leaves of the tps11 mutant plant and attenuated defense against GPA. Trehalose application restored resistance in the tps11 mutant, confirming that the lack of trehalose accumulation is associated with the inability of the tps11 mutant to control GPA infestation. Resistance against GPA was also higher in the trehalose hyper-accumulating tre1 mutant and in bacterial otsB gene-expressing plants, further supporting the conclusion that trehalose plays a role in Arabidopsis defense against GPA. Evidence presented here indicates that TPS11-dependent trehalose regulates expression of the PHYTOALEXIN DEFICIENT4 gene, which is a key modulator of defenses against GPA. TPS11 also promotes the re-allocation of carbon into starch at the expense of sucrose, the primary plant-derived carbon and energy source for the insect. Our results provide a framework for the signaling function of TPS11-dependent trehalose in plant stress responses, and also reveal an important contribution of starch in controlling the severity of aphid infestation.     

Olfactory response by the aphidophagous gall midge, Aphidoletes aphidimyza to honeydew from green peach aphid, Myzus persicae

Female adults of the aphidopagous gall midge, Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae), showed an olfactory response to honeydew excreted by the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae) under laboratory conditions. The response was only elicited by treatments with honeydew, whereas aphids, aphids with pepper plants or leaves, and pepper plants or leaves were not attractive to the midges. Dose‐dependent responses were observed from whole honeydew, honeydew volatiles extracted in pentane, and honeydew volatiles captured on Porapak Q®. When honeydew was eluted with three sequential pentane washes, a positive response was only observed from the midges for the first wash. Female midges laid more eggs on pepper plants infested with higher densities of M. persicae. The olfactory response of midges to honeydew is discussed with respect to prey location.     

阿维菌素对南方小花蝽抑制烟蚜的影响

在室内分别测定了阿维菌素类杀虫剂1% 9512-1乳油和虫螨克对南方小花蝽Orius similis的毒力,研究了无阿维菌素介入后南方小花蝽对烟蚜Myzus persicae捕食效应所受的影响,结果表明：阿维菌素类杀虫剂对南方小花蝽具有一定毒力,不同虫态对阿维菌素的敏感性有差异,卵的敏感性明显低于若虫和成虫;不同的阿维菌素类药剂对南方小花蝽的毒力存在差异,9512-1对南方小花蝽卵、若虫及成虫的LC_５０依次为11.2743±3.8613、2.6584±1.2139、2.4508±1.3266 mg·L^-１,虫螨克则为10.8699±3.7653、6.4422±2.8596、6.4738±3.0124 mg·L^-１;阿维菌素显著降低了南方小花蝽对烟蚜的捕食量和捕食能力,也降低了同种个体间的竞争能力;阿维菌素对南方小花蝽若虫的发育速率有显著的抑制作用,但对发育速率与取食量的线性关系无明显影响,而且对存活个体成虫期的产卵量无显著影响。     

Myzus persicae (green peach aphid) salivary components induce defence responses in Arabidopsis thaliana

Myzus persicae (green peach aphid) feeding on Arabidopsis thaliana induces a defence response, quantified as reduced aphid progeny production, in infested leaves but not in other parts of the plant. Similarly, infiltration of aphid saliva into Arabidopsis leaves causes only a local increase in aphid resistance. Further characterization of the defence-eliciting salivary components indicates that Arabidopsis recognizes a proteinaceous elicitor with a size between 3 and 10 kD. Genetic analysis using well-characterized Arabidopsis mutants shows that saliva-induced resistance against M. persicae is independent of the known defence signalling pathways involving salicylic acid, jasmonate and ethylene. Among 78 Arabidopsis genes that were induced by aphid saliva infiltration, 52 had been identified previously as aphid-induced, but few are responsive to the well-known plant defence signalling molecules salicylic acid and jasmonate. Quantitative PCR analyses confirm expression of saliva-induced genes. In particular, expression of a set of O -methyltransferases, which may be involved in the synthesis of aphid-repellent glucosinolates, was significantly up-regulated by both M. persicae feeding and treatment with aphid saliva. However, this did not correlate with increased production of 4-methoxyindol-3-ylmethylglucosinolate, suggesting that aphid salivary components trigger an Arabidopsis defence response that is independent of this aphid-deterrent glucosinolate.