寄主植物-蚜虫-天敌三重营养关系的化学生态学研究进展

综述了寄主植物－蚜虫－天敌三重营养关系的化学生态学研究,重点阐述了3个研究热点：①植物挥发性物质在蚜虫及其天敌选择寄主行为过程中的作用;②蚜虫信息素和蜜露对蚜虫天敌寄主选择行为的影响;③植物挥发性物质对蚜虫信息系作用的影响。对寄主植物－蚜虫－天敌三重营养关系的全面了解,将为蚜虫的综合治疗提供新思维。     

Food web structure of aphids and their parasitoids in Belgian fruit agroecosystems

Community structures of aphids and their parasitoids were studied in fruit crop habitats of eastern Belgium in 2014 and 2015. Quantitative food webs of these insects were constructed separately for each year, and divided into subwebs on three host‐plant categories, fruit crop plants, non‐crop woody and shrub plants and non‐crop herbaceous plants. The webs were analyzed using the standard food web statistics designed for binary data. During the whole study period, 78 plant species were recorded as host plants of 71 aphid species, from which 48 parasitoid species emerged. The community structure, aphid / parasitoid species‐richness ratio and trophic link number varied between the two years, whereas the realized connectance between parasitoids and aphids was relatively constant. A new plant–aphid–parasitoid association for Europe was recorded. Dominant parasitoid species in the study sites were Ephedrus persicae, Binodoxys angelicae and Praon volucre: the first species was frequently observed on non‐crop trees and shrubs, but the other two on non‐crop herbaceous plants. The potential influence, through indirect interactions, of parasitoids on aphid communities was assessed with quantitative parasitoid‐overlap diagrams. Symmetrical links were uncommon, and abundant aphid species seemed to have large indirect effects on less abundant species. These results show that trophic indirect interactions through parasitoids may govern aphid populations in fruit crop habitats with various non‐crop plants, implying the importance for landscape management and biological control of aphid pests in fruit agroecosystems.     

How aphids find their host plants,and how they don't

There is a considerable interest in trying to understand how aphids find their host plants because they are a major cause of economic losses in agricultural and horticultural production systems. Indeed, the specific behavioural sequences during host finding by aphids are one of the main reasons for their prominent role as vectors of plant pathogenic viruses. This paper reviews the visual and olfactory stimuli involved in host‐finding behaviour of aphids, both basic and applied aspects are covered. Although controlling aphids by manipulation of visual and olfactory stimuli involved in host finding and host selection can be highly successful, as shown by research, most of these measures can still be further optimised and adoption in practice is currently limited. Research therefore needs to address some critical open questions, including (a) the effects of visual contrasts on aphid behaviour; (b) the specific responses of aphids to ultraviolet (UV) light during different stages in host finding; (c) the quantification of behavioural sequences in the field and (d) the response of aphid behaviour to plant diversity at varying spatial scales; further, (e) a much more comprehensive coverage of aphid taxa and aphid ecological groups is needed in host‐finding related research to uncover ecological principles underlying this critical behaviour.     

A facultative mutualism between aphids and an invasive ant increases plant reproduction

1. The consequences to plants of ant–aphid mutualisms, particularly those involving invasive ants, are poorly studied. Ant–aphid mutualisms may increase or decrease plant fitness depending on the relative cost of herbivory by ant‐tended aphids versus the relative benefit of increased ant suppression of other (non‐aphid) herbivores. 2. We conducted field and greenhouse experiments in which we manipulated the presence and absence of cotton aphids (Aphis gossypii) on cotton plants to test the hypothesis that a mutualism between cotton aphids and an invasive ant, the red imported fire ant (Solenopsis invicta), benefits cotton plants by increasing fire ant suppression of caterpillars. We also manipulated caterpillar abundance to test whether the benefit of the mutualism varied with caterpillar density. 3. We found that more fire ants foraged on plants with cotton aphids than on plants without cotton aphids, which resulted in a significant reduction in caterpillar survival and caterpillar herbivory of leaves, flower buds, and bolls on plants with aphids. Consequently, cotton aphids indirectly increased cotton reproduction: plants with cotton aphids produced 16% more bolls, 25% more seeds, and 10% greater seedcotton mass than plants without aphids. The indirect benefit of cotton aphids, however, varied with caterpillar density: the number of bolls per plant at harvest was 32% greater on plants with aphids than on plants without aphids at high caterpillar density, versus just 3% greater at low caterpillar density. 4. Our results highlight the potential benefit to plants that host ant–hemipteran mutualisms and provide the first experimental evidence that the consequences to plants of an ant–aphid mutualism vary at different densities of non‐aphid herbivores.     

小麦蚜虫田间调查及监测技术

小麦蚜虫是世界范围内小麦生产中一类重要害虫。针对麦蚜世代历期短、繁殖力强,具有趋光、趋化及迁飞等生物学及行为习性;在田间多呈聚集分布,且麦蚜易受寄主植物抗性、天敌、气象因素及农田生态条件等生物与非生物因素影响等发生为害特点,本文阐述了我国小麦蚜虫田间调查、监测技术及防治策略,以期为我国小麦蚜虫综合防控提供基础科学支撑。     

Alarm pheromone mediates production of winged dispersal morphs in aphids

The aphid alarm pheromone ( E )- β -farnesene (EBF) is the major example of defence communication in the insect world. Released when aphids are attacked by predators such as ladybirds or lacewing larvae, aphid alarm pheromone causes behavioural reactions such as walking or dropping off the host plant. In this paper, we show that the exposure to alarm pheromone also induces aphids to give birth to winged dispersal morphs that leave their host plants. We first demonstrate that the alarm pheromone is the only volatile compound emitted from aphid colonies under predator attack and that emission is proportional to predator activity. We then show that artificial alarm pheromone induces groups of aphids but not single individuals to produce a higher proportion of winged morphs among their offspring. Furthermore, aphids react more strongly to the frequency of pheromone release than the amount of pheromone delivered. We suggest that EBF leads to a 'pseudo crowding' effect whereby alarm pheromone perception causes increased walking behaviour in aphids resulting in an increase in the number of physical contacts between individuals, similar to what happens when aphids are crowded. As many plants also produce EBF, our finding suggests that aphids could be manipulated by plants into leaving their hosts, but they also show that the context-dependence of EBF-induced wing formation may hinder such an exploitation of intraspecific signalling by plants.     

The natural occurrence of secondary bacterial symbionts in aphids

1. Many insects host secondary bacterial symbionts that are known to have wide‐ranging effects on their hosts, from host‐plant use to resistance against natural enemies. This has been most widely studied in aphids, which have become a model system to study insect–bacteria interactions. 2. While there is an increasing understanding of the role of symbionts in aphids from controlled laboratory studies, we are only beginning to explore the impact of hosting these symbionts on eco‐evolutionary dynamics in natural systems. To date, many research groups have identified bacterial symbionts from various aphid species, providing us with a bank of literature on aphid–symbiont associations in natural populations. 3. The role of secondary symbionts in aphids is discussed, and the taxonomic and geographical distribution of symbionts among aphids are summarised, and the potential reasons for the patterns observed. The need to test for multiple symbiont species (and co‐infections) across many individuals and the whole distribution range of an aphid is highlighted, including sampling on all known host‐plant species. 4. It is further important also to consider variation within the symbiont, the aphid‐host and the surrounding community, e.g. host‐plants or the natural enemies, to understand how these have the potential to mediate aphid–symbiont interactions. 5. Finally, the knowledge gained from experimental work should now be used to understand the role of aphid secondary symbionts in field systems, to fully understand the potentially far‐reaching consequences of aphid endosymbionts on community and ecosystem processes.     

The interaction of soybean aphids and soybean cyst nematodes on selected resistant and susceptible soybean lines

Soybean aphids, Aphis glycines Matsumura, and soybean cyst nematodes, Heterodera glycines Ichinohe, are economic pests of soybean, Glycine max (L.) Merr., in the north‐central United States. Combined, these pests may account for 20–50% of yield reductions in a soybean crop. Only limited information is available concerning the interaction of these two pests on soybean production. During the summers of 2006 and 2007, we conducted a field‐experiment near Urbana, IL, to evaluate the effect of resistant and susceptible soybean lines on the development and reproduction of both pests in combination with each other. We also examined how each pest, as well as their interaction, affected the yield of susceptible and resistant soybean lines. Soybean plants grown within caged plots were infested with soybean aphids and soybean cyst nematodes; cumulative aphid days and soybean cyst nematode egg densities were determined at the end of each growing season. Soybean aphids were able to survive on all four soybean lines in both years of this study; however, aphid‐resistant lines generally had fewer cumulative aphid days than aphid‐susceptible lines. Likewise, nematode‐resistant lines typically had fewer eggs than nematode‐susceptible lines. During both years, we failed to observe a significant interaction between these two pests on the reproduction of one another. Yield data from 2006 was inconclusive; however, results from 2007 suggest that yield‐loss when soybean aphids and soybean cyst nematodes occur jointly is not significantly greater than when these two pests occur independently. The relationship between these two pests, and our inability to observe an interaction, are discussed.     

Effect of a fungal lectin from Xerocomus chrysenteron (XCL) on the biological parameters of aphids

Aphids are important pests of crop plants in Europe. Increasing resistance of aphids to insecticides and their side effects on the environment and non target organism's including human's stimulated research on alterative methods of aphid control, including the use of entomotoxic proteins. Lectins are carbohydrate binding proteins that are widely distributed in nature; they have been isolated from microorganisms, fungi, plants and animals. Several of these proteins were tested for their potential biocide effect on plenty of pests. A fungal lectin, namely Xerocomus Chrysenteron lectin (XCL) was previously purified and was shown to be toxic for several pests including aphids. XCL was clearly the most toxic lectin against M. persicae. In this work, bioassays using artificial diets incorporating a broad range of XCL concentrations (from 10 microg x ml(-1) to 5000 microg x ml(-1)) were developed to assess the negative effects of XCL on the biological parameters (development duration, weight and fecundity) of M. persicae a polyphagous aphid found on more than 400 host plant species and transmitting more than 100 viral diseases. A significant mortality of aphids was observed, corresponding to the LC50 and LC90 of 0, 46 and 6, 02 mg/ml respectively after 24hrs. Significant differences of M. persicae weight, development duration and fecundity (P < 0.05) was observed between the tested XCL concentrations. Conavalia ensifomris lectin (ConA) was included as lectin reference on the bioassay experiments and was shown to be less toxic and induced lower negative changes in M. persicae biological parameters when compared with XCL.     

Attachment forces of pea aphids (Acyrthosiphon pisum) on different legume species

1. Sympatric populations of insects adapted to different host plants are good model systems not only to study how they adapt to the chemistry of their food plant, but also to investigate whether morphological modifications evolved enabling them to live successfully on a certain plant species. 2. The pea aphid, Acyrthosiphon pisum (Harris) encompasses at least 11 genetically distinct sympatric host races, each showing a preference for a certain legume species. The leaflet surfaces of these legumes differ considerably in their wax coverage. 3. It was investigated whether the attachment structures of three pea aphid genotypes from different host races are adapted to the different surface properties of their host plants and whether they show differences in their attachment ability on the respective host and non‐host plants. 4. The surface morphology of plants and aphid tarsi was examined using SEM (scanning electron microscopy). The ability of the aphids to walk on specific surfaces was tested using traction force measurements. 5. The presence of wax blooms on the leaflets lowers the aphids' attachment ability considerably and diminishes their subsequent attachment on ‘neutral’ surfaces like glass. The pea aphid host races differ in their ability to walk on certain surfaces. However, the genotype from the adapted aphid host race was not necessarily the one with the best walking performance on their host plant. All aphids, regardless of the original host plant, were most efficient on the neutral control surface glass. The general host plant Vicia faba was the plant with the most favourable surface for all aphid host races.     

蚜虫的表型可塑性及其遗传基础

表型可塑性(phenotypic plasticity)是有机体在适应生物或非生物环境时呈现不同表型的能力,并且有遗传基础。蚜虫是农林业的重要经济害虫,易受外部环境因素和自身遗传因素的影响而表现出表型的可塑性。本文综述了外部环境因素（如寄主植物、温度、光照、天敌等）的变异对蚜虫表型的影响。总体来说,蚜虫表型会因寄主植物的种类、品系以及发育阶段和营养状况的不同而有所差异; 温度变化对不同蚜虫种类的生殖力、生存力以及有翅蚜产生与否有极大影响。研究人员利用RAPD-PCR、微卫星等分子遗传标记确认寄主植物和温度是造成蚜虫种群遗传分化的重要因素。就内部因素而言,不同的蚜虫种类以及同一种蚜虫的不同克隆系在表型和遗传进化上也有不同程度的差异,在蚜虫受外界条件影响的不同虫态以及不同体色克隆系、不同生活周期的类群之间, 其生物学、生态学和遗传学都有所差异。分析上述各个因素对蚜虫表型可塑性的影响,对于蚜虫的生态进化研究和有效治理蚜害均有重要意义。本文在最后讨论了还有必要深入研究的诸多问题,如表观遗传调控,包括DNA甲基化、基因所在的核小体上的组蛋白的共价修饰和染色质重塑、siRNA介导的基因沉默以及微RNA（microRNA 或 miRNA）调控的基因表达变化等,又如有翅蚜的表型和遗传学研究,以及全球气候变化对蚜虫的生态进化的影响等问题。     

Plant-aphid interactions: molecular and ecological perspectives

Many aphids are major agricultural pests because of their unparalleled reproductive capacity and their ability to manipulate host plant physiology. Aphid population growth and its impact on plant fitness are strongly influenced by interactions with other organisms, including plant pathogens, endophytes, aphid endosymbionts, predators, parasitoids, ants, and other herbivores. Numerous molecular and genomic resources have recently been developed to identify sources of aphid resistance in plants, as well as potentially novel targets for control in aphids. Moreover, the same model systems that are used to explore direct molecular interactions between plants and aphids can be utilized to study the ecological context in which they occur.     

Evolutionary ecology of the interactions between aphids and their parasitoids

Many organisms, including entomopathogenous fungi, predators or parasites, use aphids as ressources. Parasites of aphids are mostly endoparasitoid insects, i.e. insects which lay eggs inside the body of an other insect which will die as a result of their development. In this article, we review the consequences of the numerous pecularities of aphid biology and ecology for their endoparasitoids, notably the Aphidiinae (Hymenoptera: Braconidae). We first examine the various mechanisms used by aphids for defence against these enemies. We then explore the strategies used by aphidiine parasitoids to exploit their aphid hosts. Finally, we consider the responses of both aphids and parasitoids to ecological constraints induced by seasonal cycles and to environmental variations linked to host plants and climate. The fundamental and applied interest of studying these organisms is discussed.     

Field‐level effects of preventative management tactics on soybean aphids (Aphis glycines Matsumura) and their predators

A 2‐year field experiment was conducted in northern Illinois to evaluate the effects of host plant resistance and an insecticidal seed treatment (thiamethoxam) on soybean aphids, Aphis glycines Matsumura and their predators. Densities of soybean aphids varied between the 2 years of the experiment. During both years, resistant plants experienced fewer cumulative aphid days than susceptible plants. Populations of soybean aphids on resistant plants rarely exceeded the economic injury level of 250 soybean aphids per plant. The use of thiamethoxam reduced cumulative aphid days in 2007, but not in 2008. Although soybean aphids reached densities that were sufficient to cause yield‐loss for untreated and susceptible plants, no yield‐benefit was associated with using the two management tactics in either year. This latter finding suggests that densities of soybean aphids need to be greater and sustained for a longer period of time than what we observed if the two management tactics are expected to provide a yield‐benefit. Monitoring natural enemies revealed that densities of key aphidophagous predators were relatively unaffected by host plant resistance or thiamethoxam; the effect of these management tactics on densities of predators, as well as the effectiveness of the method used to sample predators, is discussed.     

How aphid alarm pheromone can control aphids: a review

Aphids are the major pests of arable crops, mostly in temperate regions. They are monophagous as well as polyphagous. They inflict damage in brassica, potato, cotton, vegetable and fruit crops. They damage their host plant directly by feeding upon their phloem sap, or indirectly by transmitting pathogens to them. Their life cycle can be autoecious as well as heteroecious. Aphids use semiochemicals for various purposes, in gathering information from their environment and for communication among themselves. They protect themselves from predators and parasitoids by escape response which is arbitrated by use of alarm pheromone signalling. When alarm pheromone, (E)-ß-farnesene, is released, nearby aphids exhibit a variety of behaviours like moving away, running, dropping off the plant and even attacking the predator. Previous studies of integrated pest management strategies have been aimed at the usage of alarm pheromone. However, scientists require complete knowledge of aphid ecology as well as aphid interaction with its natural enemies to establish efficient and viable biological control. This review presents analysis of the existing aphid pest management methodologies and effectiveness of alarm pheromone on aphids and their natural enemies.     

药剂包衣对苗期大豆蚜防治效果与安全性评价

【目的】大豆蚜Aphis glycines(Matsumura)是大豆上最重要的害虫之一。传统控制大豆蚜虫仍然以达到防治指标时大量喷洒化学药剂为主,危害人畜和环境安全。只有在大豆蚜发生初期进行有效防控,使其田间种群不能及时顺利的建立,从而实现无公害绿色防控。【方法】对筛选出的3种内吸式杀虫剂按不同浓度拌种包衣进行大田小区试验,调查分析包衣处理对大豆蚜、天敌以及大豆田其他害虫的影响和控制作用,同时对包衣处理后的大豆安全性、产量和品质进行评估。【结果】药剂拌种包衣处理能够显著压低苗期大豆蚜虫口基数,2014年对照区与处理区蚜量最高峰值比值最大达到448.15;同时对苗期大豆田间的双斑萤叶甲Monolepta hieroglyphica(Motschulsky)有很好的控制作用,处理区与对照区的受害株率差异极显著;并且保护了自然天敌种群;药剂拌种包衣处理在显著增产的同时还有效提升了大豆品质;经权威检测,收获后的籽粒在检出限内无药剂残留。【结论】药剂拌种包衣处理能有效控制苗期大豆蚜,不杀伤天敌,安全、无毒、无残留,而且增产显著,是比较理想的轻简无公害防控手段。     

Nutritional enhancement of host plants by aphids - a comparison of three aphid species on grasses

Three aphid species were compared with respect to ability of enhancing the nutritional quality of their host plants. Rhopalosiphum padi, which does not induce macroscopic changes in its host plants, was compared with Schizaphis graminum and Diuraphis noxia, both of which induce distinctive types of chlorotic lesions. Phloem sap samples were collected from severed stylets of feeding aphids and from exudates of cut leaves of plants uninfested or infested with each aphid species. Samples were analyzed for concentrations of individual amino acids.Compared to R. padi, S. graminum ingested phloem sap with a two-fold higher concentration of amino acids and a much higher proportion of essential amino acids. Similar differences between these two aphid species were observed on both wheat and barley. For each aphid species, the absolute concentrations of amino acids and the relative proportions of essential amino acids were similar between the two host plants. Effects of D. noxia were similar to those of S. graminum, though less dramatic. Exudates from leaves infested with each aphid species showed relative concentrations of individual amino acids that were similar to those in the corresponding stylet exudates. Based on comparison of stylet exudates and cut leaf exudates from infested and uninfested plants, R. padi seems to have little effect on amino acid composition of phloem. Changes in the phloem induced by both S. graminum and D. noxia appear to be systemic, affecting at least the whole leaf they are feeding on. The changes observed for D. noxia and for S. graminum are likely to be nutritionally advantageous for the aphids and are expected to affect the aphids' dependence on nutritional supplementation by intracellular symbionts (Buchnera).     

Changes in activity of superoxide dismutase and catalase within cereal aphids in response to plant o-dihydroxyphenols

Abstract:  Superoxide dismutase and catalase activity was found in the bird cherry-oat aphid, Rhopalosiphum padi (L.) and the grain aphid, Sitobion avenae (F.). Among the aphid morphs studied, the highest activity of the antioxidant enzymes was noted for winged adults (alatae) and the lowest for wingless (apterae) ones. Higher activity of superoxide dismutase and catalase was observed in the polyphagous species R. padi that alternates between woody host plants and grasses. On some ocassions, activity of superoxide dismutase in cereal aphids was increased by twofold, when aphids were exposed to toxic plant o -dihydroxyphenols. An opposite tendency was observed in case of activity of the catalase that was strongly reduced within body of phenolics-treated insects. Among the plant allelochemicals studied, caffeic acid showed the strongest effect on the activity of the antioxidant enzymes of the cereal aphids. The experiments carried out indicate that antioxidant enzymes might play an important role in interactions between cereal aphids and their host plants.     

Ants indirectly reduce the reproductive performance of a leafless shrub by benefiting aphids through predator deterrence

Ant–aphid mutualisms can generate cascade effects on the host plants, but these impacts depend on the ecological context. We studied the consequences of ant–aphid interactions on the reproductive performance of a Mediterranean leafless shrub (Retama sphaerocarpa), through direct and indirect effects on the arthropod community. By manipulating the presence of ants and aphids in the field, we found that ants increased aphid abundance and their persistence on the plant and reduced aphid predators by nearly half. However, the presence of ants did not affect the abundance of other plant herbivores, which were relatively scarce in the studied plants. Aphids, and particularly those tended by ants, had a negative impact on the plant reproductive performance by significantly reducing the number of fruits produced. However, fruit and seed traits were not changed by the presence of aphids or those tended by ants. We show that ants favoured aphids by protecting them from their natural enemies but did not indirectly benefit plants through herbivory suppression, resulting in a net negative impact on the plant reproductive performance. Our study suggests that the benefits obtained by plants from hosting ant–aphid mutualisms are dependent on the arthropod community and plant traits.

     

Metabolic Engineering of Plant-derived (E)-β-farnesene Synthase Genes for a Novel Type of Aphid-resistant Genetically Modified Crop Plants

Aphids are major agricultural pests that cause significant yield losses of crop plants each year.Excessive dependence on insec-ticides for long-term aphid control is undesirable because of the development of insecticide resistance,the potential negative effects on non-target organisms and environmental pollution.Transgenic crops engineered for resistance to aphids via a non-toxic mode of action could be an efficient alternative strategy.(E)-β-Farnesene (EβF) synthases catalyze the formation of EβF,which for many pest aphids is the main component of the alarm pheromone involved in the chemical communication within these species.EβF can also be synthesized by certain plants but is then normally contaminated with inhibitory compounds.Engineering of crop plants capable of synthesizing and emitting EβF could cause repulsion of aphids and also the attraction of natural enemies that use EβF as a foraging cue,thus minimizing aphid infestation.In this review,the effects of aphids on host plants,plants' defenses against aphid herbivory and the recruitment of natural enemies for aphid control in an agricultural setting are briefly introduced.Furthermore,the plant-derived EβF synthase genes cloned to date along with their potential roles in generating novel aphid resistance via genetically modified approaches are discussed.