蚜虫寄主专化型及其成因

综述了蚜虫寄主专化型产生的可能原因。蚜虫寄主专化型的形成在生态学水平上主要与蚜虫对寄主的选择识别能力、天敌作用、其他共生（共存）生物作用、抗药性等有关;并且具有一定的遗传基础,主要表现在蚜虫体内酶系的变化、染色体变异、有性繁殖中同型交配行为及种群的遗传分化上。但是对某种寄主专化型蚜虫而言,其具体的成因尚不明确。     

棉蚜寄主专化型及其形成的行为机理

通过生活在甜瓜和棉花上的棉蚜Aphisgossypii Glover的行为,研究棉蚜的寄主专化型及其形成的行为机理。生物学观察显示： 两类棉蚜在寄主植物相互交换以后,定居数显著减少,棉花蚜型棉蚜的繁殖系数及若虫存活率显著下降,说明棉蚜存在甜瓜蚜型和棉花蚜型两种寄主专化型。通过刺探电位技术研究棉蚜的取食行为,以探索其寄主专化型形成的行为机理。结果表明： 甜瓜蚜型棉蚜在棉花上的取食行为容易被中断,但其口针定位韧皮部的能力并没有显著削弱;而棉花蚜型棉蚜在甜瓜上的取食行为受到更大的影响,口针无法顺利定位至韧皮部,并在2 h内根本无法在筛管内取食。生物学观察和EPG取食行为分析都显示： 与甜瓜蚜型棉蚜相比,棉花蚜型棉蚜对寄主的要求更严格-寄主专化程度更高,对寄主的利用率更高。     

Combined effects of host-plant resistance and intraguild predation on the soybean aphid parasitoid Binodoxys communis in the field

Soybean varieties that exhibit resistance to the soybean aphid Aphis glycines have been developed for use in North America. In principle, host-plant resistance to soybean aphid can influence the interactions between the soybean aphid and its natural enemies. Resistance could change the quality of soybean aphids as a food source, the availability of soybean aphids, or resistance traits could directly affect aphid predators and parasitoids. Here, we focus on the effect of soybean aphid resistance on the interactions between soybean aphids, the parasitoid Binodoxys communis (Hymenoptera: Braconidae), and predators of these two species. We determined whether host-plant resistance affected within-season persistence of B. communis by releasing parasitoids into resistant and susceptible soybean plots. We observed higher B. communis densities in susceptible soybean plots than in resistant plots. There were also higher overall levels of intraguild predation of B. communis in susceptible plots, although the per-capita risk of intraguild predation of B. communis was affected neither by plant genotype nor by aphid density. We discuss these effects and whether they were caused by direct effects of the resistant plants on B. communis or indirect effects through soybean aphid or predators.     

Intraspecific variation of Myzus persicae on sugar beet (Beta vulgaris)

Differences in inherited resistance among seven sugar-beet stocks had similar effects on Myzus persicae clones representing the range of variation in aphid response to resistant and susceptible sugar beet observed in fifty-eight clones collected between 1969 and 1971. Three sugar-beet stocks were consistently resistant. Statistically significant interactions between beet stocks and aphid clones did not indicate the existence of biotypes with specific abilities to overcome resistance. M. persicae clones differed in their vigour of colonizing sugar beet, irrespective of the differences between beet stocks. The readiness of adult aphids to settle determined the size of aphid population produced and included a component related to the response of the aphid clone to sugar beet as a host, and a component related to the resistance ranking of the beet stock. Breeding sugar beet with resistance to aphids will be simplified, as the results indicate that, at present, differences between aphid biotypes need not be considered a problem.     

Host plant resistance to aphids in cultivated crops: Genetic and molecular bases,and interactions with aphid populations

Host plant resistance is an efficient and environmentally friendly means of controlling insects, including aphids, but resistant-breaking biotypes have occurred in several plant–aphid systems. Our review of the genetic and molecular bases of aphid resistance in crop species emphasizes the limited number of aphid resistance genes and alleles. Inheritance of aphid resistance may be monogenic (dominant or recessive genes) or polygenic. Two dominant, aphid resistance genes have been isolated to date. They both encode NBS-LRR proteins involved in the specific recognition of aphids. Strategies to ensure aphid resistance effectiveness and durability are discussed. Innovative research activities are proposed.     

Facultative symbionts are associated with host plant specialization in pea aphid populations

The pea aphid, Acyrthosiphon pisum, shows significant reproductive isolation and host plant specialization between populations on alfalfa and clover in New York. We examine whether specialization is seen in pea aphids in California, and whether fitness on alternative host plants is associated with the presence of bacterial symbionts. We measured the fitness of alfalfa- and clover-derived aphids on both types of plants and found no evidence for specialization when all aphid lineages were considered simultaneously. We then screened all aphids for the presence of four facultative bacterial symbionts: PAR, PASS, PABS and PAUS. Aphids with PAUS were host-plant specialized, having twice as many offspring as other aphids on clover, and dying on alfalfa. Other aphids showed no evidence of specialization. Additionally, aphids with PABS had 50% more offspring than aphids with PASS when on alfalfa. Thus, specialist and generalist aphid lineages coexist, and specialization is symbiont associated. Further work will resolve whether PAUS is directly responsible for this variation in fitness or whether PAUS is incidentally associated with host-plant specialized aphid lineages.     

Genetic mapping of three quantitative trait loci for soybean aphid resistance in PI 567324

Host-plant resistance is an effective method for controlling soybean aphid (Aphis glycines Matsumura), the most damaging insect pest of soybean (Glycine max (L.) Merr.) in North America. Recently, resistant soybean lines have been discovered and at least four aphid resistance genes (Rag1, Rag2, Rag3 and rag4) have been mapped on different soybean chromosomes. However, the evolution of new soybean aphid biotypes capable of defeating host-plant resistance conferred by most single genes demonstrates the need for finding germplasm with multigenic resistance to the aphid. This study was conducted to map quantitative trait loci (QTL) for aphid resistance in PI 567324. We identified two major QTL (QTL_13_1 and QTL_13_2) for aphid resistance on soybean chromosome 13 using 184 recombinant inbred lines from a ‘Wyandot'' × PI 567324 cross. QTL_13_1 was located close to the previously reported Rag2 gene locus, and QTL_13_2 was close to the rag4 locus. A minor QTL (QTL_6_1) was also detected on chromosome 6, where no gene for soybean aphid resistance has been reported so far. These results indicate that PI 567324 possesses oligogenic resistance to the soybean aphid. The molecular markers closely linked to the QTL reported here will be useful for development of cultivars with oligogenic resistance that are expected to provide broader and more durable resistance against soybean aphids compared with cultivars with monogenic resistance.     

Trichomes of Lycopersicon spp. and their effect on Myzus persicae (Sulzer) (Hemiptera: Aphididae)

Abstract The green peach aphid ( Myzus persicae ) is becoming resistant to many of the pesticides used in the tomato ( Lycopersicon esculentum ) industry. A potential alternative method of control is host-plant resistance, which may be mediated by trichomes. Resistance to the green peach aphid was assessed for six accessions of L. hirsutum, three of L. pennellii and a L. esculentum control. Mortality, entrapment and number of aphids unaffected by trichomes were recorded 24, 48 and 72 h after placement. Using multiple regression against counts of glandular and non-glandular trichomes, the role of foliar pubescence in resistance to the aphid was established. Lycopersicon pennellii possessed the highest level of resistance after 72 h and type IV trichome density accounted for 26.5% of variance in mortality ( P  < 0.001). Low densities of type IV and high densities of type VII trichomes increased the number of aphids 'unaffected' by trichomes, a relationship accounting for 29% of variance ( P  < 0.001). Information on which types of trichomes most strongly affect resistance to insect pests will aid future breeding and L. pennellii shows potential for use in breeding programs to increase levels of host-plant resistance in L. esculentum.     

Proteomic profiling of aphid Macrosiphum euphorbiae responses to host-plant-mediated stress induced by defoliation and water deficit

Abiotic and biotic host-plant stress, such as desiccation and herbivory, may strongly affect sap-sucking insects such as aphids via changes in plant chemicals of insect nutritional or plant defensive value. Here, we examined (i) water deprivation and (ii) defoliation by the beetle Leptinotarsa decemlineata as stresses indirectly affecting the aphid Macrosiphum euphorbiae via its host plant Solanum tuberosum. For plant-induced stress, aphids were reared on healthy vs. continuously stressed potato for 14 days (no watering; defoliation maintained at approximately 40%). Aphid performance under stress was correlated with metabolic responses monitored by profiling of the aphid proteome. M. euphorbiae was strongly affected by water stress, as adult survival, total aphid number and biomass were reduced by 67%, 64%, and 79%, respectively. Aphids performed normally on defoliated potato, indicating that they were unaffected or able to compensate any stress induced by plant defoliation. Stressed aphid proteomes revealed 419-453 protein spots, including 27 that were modulated specifically or jointly under each kind of host-plant stress. Reduced aphid fitness on water-stressed plants mostly correlated with modulation of proteins involved in energy metabolism, apparently to conserve energy in order to prioritize survival. Despite normal performance, several aphid proteins that are known to be implicated in cell communication were modulated on defoliated plants, possibly suggesting modified aphid behaviour. The GroEL protein (or symbionin) of the endosymbiont Buchnera aphidicola was predominant under all conditions in M. euphorbiae. Its expression level was not significantly affected by aphid host-plant stresses, which is consistent with the high priority of symbiosis in stressed aphids.     

棉花型和黄瓜型棉蚜对寄主植物的适合度

采用寄主转换建立生命表及触角电位 (EAG)方法比较了棉蚜两寄主专化型 (棉花型和黄瓜型 )对不同夏季作物的适合度。结果表明 ,棉花型蚜转到黄瓜上及黄瓜型蚜转接到棉花上均不能正常产仔繁殖和建立种群。两种蚜型均不能在茄子和苋菜上建立种群。黄瓜型蚜能在豇豆上建立种群 ,但棉花型蚜不能 ,表现出两种寄主型棉蚜对夏寄主利用上存在显著差异。受棉蚜为害 12～ 36 h后的黄瓜或棉花植株仍不适合于棉花型或黄瓜型蚜 ,表现出黄瓜型蚜不能在被棉花型蚜为害过的棉株上正常存活和繁殖 ,棉花型蚜也不能在被黄瓜型蚜为害过的黄瓜苗上存活和繁殖。两种寄主型蚜对不同寄主叶片丙酮提取物的触角电位表明 ,黄瓜型蚜对棉花、哈密瓜和马铃薯叶片提取物的反应显著强于棉花型蚜 ,而对黄瓜和甜瓜叶片提取物的反应上两种蚜型差异不显著。文中同时对棉花型和黄瓜型棉蚜产生的生态机制进行了讨论。     

Discovery of English grain aphid (Hemiptera: Aphididae) biotypes in China

The English grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae), is an important pest insect of wheat, Triticum aestivum (L.), in China. Grain aphid biotypes are necessary to breed aphid-resistant wheat varieties; however, none have currently been identified. Here, we describe a method to identify grain aphid biotypes and survey the aphid biotype variation in the wheat growth area of China. Clones of S. avenae were collected from 11 locations in China and used to establish culture populations. These populations were then used to assess the resistance of 12 wheat varieties. Based on resistance responses, seven differential hosts were selected to identify the biotype of S. avenae: Amigo, 'Fengchan No. 3', Zhong 4 wumang, JP1, L1, 885479-2, and 'Xiaobaidongmai'. S. avenae was ultimately classified into five biotypes: EGA I, EGA II, EGA III, EGA IV, and EGA V. These methods provide a mechanism to detect the variation and evolution of grain aphids in different wheat-growing locations and also allow for selection of appropriate aphid-resistant germplasm for wheat breeding of commercial wheat cultivars.     

A comparison of maternal effects and current environment on vital rates of Aphis nerii, the milkweed–oleander aphid

Abstract.  1. Non-Mendelian maternal effects, the effects of maternal phenotype or environment on offspring phenotype, have been documented in numerous taxa. By affecting offspring vital rates (birth, death, and movement), maternal effects have the potential to influence population dynamics. However, relatively few studies have directly linked maternal phenotype or environment to offspring vital rates. Additionally, even fewer studies have compared the magnitude of across-generation effects (i.e. maternal effects) to within-generation effects.
2. Because of their telescoping generations, aphids can be strongly influenced by maternal effects. The effects of maternal density and maternal host-plant species on offspring survival, fecundity, and alate formation were investigated experimentally in Aphis nerii , the milkweed–oleander aphid.
3. Additionally, the relative strength of maternal effects were compared with those operating within a generation. Therefore, in another set of experiments, the effects of current density and host-plant species (within-generation effects) on aphid vital rates were examined.
4. While maternal effects were present, within-generation effects were much stronger and more strongly influenced aphid vital rates. Within a generation, aphids exhibited density-dependent survival, fecundity, and alate formation and these effects varied among host-plant species.
5. These results indicate that while maternal effects have the potential to affect population dynamics, this potential is not always met. Additionally, the current environment, not the environment of previous generations, more strongly impacts population dynamics.     

Proteomic analysis of secreted saliva from Russian Wheat Aphid (Diuraphis noxia Kurd.) biotypes that differ in virulence to wheat

Diuraphis noxia, Russian Wheat Aphid (RWA), biotypes are classified by their differential virulence to wheat varieties containing resistance genes. RWA salivary proteins, unlike those of most aphid species, cause foliar damage and physiological alterations in plants. A comparative proteomic analysis of secreted saliva from four differentially virulent RWA biotypes identified thirty-four individual proteins. The five major proteins were glucose dehydrogenase, lipophorin, chitinase, CiV16.8g1-like, and lava lamp. Fourteen proteins quantitatively varied among biotypes; trehalase, β-N-acetylglucosaminidase (chitinase), two separate glucose dehydrogenases, calreticulin, aminopeptidase, acetylglucosaminyltransferase, hydroxymethylglutaryl-CoA lyase, acyltransferase, ficolin-3, lava lamp, retinaldehyde-binding protein, and two proteins of unknown function. Fifty-four percent of spectral counts were associated with glucose dehydrogenase, which is thought to detoxify plant defensive compounds. One-dimensional electrophoresis detected nine protein bands from 9 to 60 kDa that quantitatively differed. Two-dimensional electrophoresis identified six major gel zones with quantitative and qualitative variance in proteins. Our findings reveal that the salivary proteome of RWA, a phytotoxic aphid, differs considerably from those reported for nonphytotoxic aphids. The potential roles of proteins used in the general plant feeding processes of aphids and those that are potential phytotoxins related to aphid virulence are discussed.     

紫外辐射对两种体色型麦长管蚜生物学特征的影响

紫外辐射对蚜虫产生了强烈的选择压力,可导致蚜虫种下遗传分化,而紫外辐射对蚜虫种下体色分化的影响尚不清楚.本研究选定两种不同体色型的麦长管蚜初生若蚜经过30 W紫外线（UVB波）照射5 d,每天30 min,在不同温度（15 ℃、20 ℃和25 ℃）和不同小麦品种（小偃-22、Astron）条件下,通过在人工智能控制温室内单头饲养,分别测定了各处理麦长管蚜的发育历期、体质量差及相对日均体质量增长率等生物学参数.结果表明：在品种小偃-22上,温度较低时,紫外辐射显著延缓了绿色型麦长管蚜生长,而温度较高时,紫外辐射显著延缓了红色型麦长管蚜生长;在品种Astron上,温度较低时,紫外辐射显著延缓了绿色型麦长管蚜生长,而温度较高时,紫外辐射对两种体色型麦长管蚜生长均无显著影响.说明两种体色型蚜虫对紫外胁迫的反应不同,且紫外胁迫对麦长管蚜的影响与温度和小麦品种有关.     

Host preference of plant genotypes is altered by intraspecific competition in a phytophagous insect

Variation among aphid genotypes leads them to preferentially colonize different host-plant genotypes. In a natural community, different genotypes within a species are expected to coexist on a single host plant, and these aphids can interact, potentially, altering host-plant preferences. Using a model aphid (Sitobion avenae) and barley (Hordeum vulgare) system, we compared aphid preference and performance in one- or two-genotype colonies in pots with genetically diverse host plants (6 genotypes) or genetically uniform host plants (1 genotype per pot). Aphid host preference was shown to differ when a second aphid genotype was present, with one aphid genotype exhibiting a preference change due to the genotypic identity of the second aphid. The population growth rate of the aphids was not influenced by the competitor, and thus, we conclude that these effects are due to aphid distribution (preference) rather than effects through performance. Our work demonstrates that within a complex ecological community, an individual’s behavior can be influenced by interactions with other genotypes within the same species, as well as interactions with genotypes of other species.     

Host range expansion of an introduced insect pest through multiple colonizations of specialized clones

Asexuality confers demographic advantages to invasive taxa, but generally limits adaptive potential for colonizing of new habitats. Therefore, pre-existing adaptations and habitat tolerance are essential in the success of asexual invaders. We investigated these key factors of invasiveness by assessing reproductive modes and host-plant adaptations in the pea aphid, Acyrthosiphon pisum, a pest recently introduced into Chile. The pea aphid encompasses lineages differing in their reproductive mode, ranging from obligatory cyclical parthenogenesis to fully asexual reproduction. This species also shows variation in host use, with distinct biotypes specialized on different species of legumes as well as more polyphagous populations. In central Chile, microsatellite genotyping of pea aphids sampled on five crops and wild legumes revealed three main clonal genotypes, which showed striking associations with particular host plants rather than sampling locations. Phenotypic analyses confirmed their strong host specialization and demonstrated parthenogenesis as their sole reproductive mode. The genetic relatedness of these clonal genotypes with corresponding host-specialized populations from the Old World indicated that each clone descended from a particular Eurasian biotype, which involved at least three successful introduction events followed by spread on different crops. This study illustrates that multiple introductions of highly specialized clones, rather than local evolution in resource use and/or selection of generalist genotypes, can explain the demographic success of a strictly asexual invader.     

Factors affecting transmission of fungal pathogens of aphids

Fungal pathogens are the most important pathogens of aphids (Homoptera: Aphididae), and epizootics, particularly those caused by Entomophthorales (Zygomycota), are frequently observed and often rapidly reduce aphid populations. Fungi in the Hypocreales (Ascomycota) are less commonly found infecting aphids but can be important. The transmission of aphid fungal pathogens is affected by many factors, including: host biology and structure, pathogen characteristics, host-plant characteristics, and environmental factors. This paper is an overview of selected factors affecting transmission of aphid pathogens.     

Chemical defence,offence and alliance in ants–aphids–ladybirds relationships

Chemicals, which mediate the interactions between aphids, ladybirds and ants, are reviewed. Special emphasis is laid on autogenous and plant-derived chemical defence in aphids and ladybirds. Evidences for chemical cues used during foraging and oviposition in ladybirds are assessed. Possible mutualistic interaction between plants and the third trophic level is illustrated by the as yet few reports of indirect plant-defence volatiles induced by aphids or coccids attracting parasitoids or ladybirds. The use of chemical signals allowing aphid parasitoids or ladybirds to squeeze into ant–aphid mutualistic association is briefly described. Questions are raised and hypotheses suggested which could stimulate further research on aphid host-plant influence on ladybird foraging behaviour and fitness, and on the cues used by aphid-web partners for their mutual recognition.     

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.     

Field and laboratory evaluations of soybean lines against soybean aphid (Hemiptera: Aphididae)

The soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), is a major pest of soybean, Glycine max (L.). Merr., that significantly reduces yield in northern production areas of North America. Insecticides are widely used to control soybean aphid outbreaks, but efforts are underway to develop host plant resistance as an effective alternative management strategy. Here, previously identified resistant lines were evaluated in laboratory tests against field-collected populations of soybean aphid and in field-plot tests over 2 yr in South Dakota. Six lines previously identified with resistance to soybean aphid--Jackson, Dowling, K1639, Cobb, Palmetto and Sennari--were resistant in this study, but relatively high aphid counts on Tie-feng 8 in field plots contrasted with its previously reported resistance. Bhart-PI 165989 showed resistance in one of two laboratory tests, but it had relatively large aphid infestations in both years of field tests. Intermediate levels of soybean aphid occurred in field plots on lines previously shown to have strong (Sugao Zairai, PI 230977, and D75-10169) or moderate resistance to soybean aphid (G93-9223, Bragg, Braxton, and Tracy-M). Sugao Zairai also failed to have a significant proportion of resistant plants in two laboratory tests against aphids field-collected in 2008, but it was resistant in laboratory tests with aphids collected in 2002, 2005, and 2006. Overall, results showed that lines with Rag (i.e., Jackson) or Rag1 gene (i.e., Dowling) had low aphid numbers, whereas lines with Rag2 (i.e., Sugao Zairai, Sennari) had mixed results. Collectively, responses of soybean aphid populations in laboratory and field tests in 2008 resembled a virulence pattern reported previously for biotype 3 soybean aphids, but virulence in soybean aphid populations was variable and dynamic over years of the study. These results, coupled with previous reports of biotypes virulent to Rag1, suggest that deployment of lines with a single aphid-resistance gene is limited for soybean aphid management, and that deployment strategies relying on multiple resistance genes may be needed to effectively use plant resistance against soybean aphid.