Specific herbivore-induced volatiles defend plants and determine insect community composition in the field

In response to insect attack, plants release complex blends of volatile compounds. These volatiles serve as foraging cues for herbivores, predators and parasitoids, leading to plant-mediated interactions within and between trophic levels. Hence, plant volatiles may be important determinants of insect community composition. To test this, we created rice lines that are impaired in the emission of two major signals, S-linalool and (E)-β-caryophyllene. We found that inducible S-linalool attracted predators and parasitoids as well as chewing herbivores, but repelled the rice brown planthopper Nilaparvata lugens, a major pest. The constitutively produced (E)-β-caryophyllene on the other hand attracted both parasitoids and planthoppers, resulting in an increased herbivore load. Thus, silencing either signal resulted in specific insect assemblages in the field, highlighting the importance of plant volatiles in determining insect community structures. Moreover, the results imply that the manipulation of volatile emissions in crops has great potential for the control of pest populations.     

The constraints of selecting for insect resistance in plantation trees

• 1 High productivity in plantations of exotic tree species is achieved by management for fast growth in the absence of the full complex of co‐evolved insect herbivores. In the case of Eucalyptus, silvicultural selection for desirable wood traits is concomitant with a trade‐off against defence and a reduction of chemical and genetic diversity. These factors, combined with accidental introductions, rapid insect evolution and the emergence of new pests, increase the likelihood that future plantations will need insect pest management to maintain productivity.
• 2 Forestry researchers have suggested that selecting for resistant genotypes may be beneficial in insect control. There are, however, significant differences between long‐lived trees and annual crops that make this approach unlikely to be successful. This is illustrated using several examples of research into resistance to insect herbivores in trees.
• 3 Selection for resistance to insects in trees requires an assessment of trial plantations for heritable variation in insect damage and then a determination of the effect of variation in resistance on insect population parameters. Identifying rare resistant genotypes using markers is difficult because many factors interact to produce a resistant phenotype, and phytophagous insects have less intimate relationships with their host than pathogens, resulting in weak associations with genetic loci.
• 4 If resistant genotypes are identified, their widespread deployment in plantations might not provide satisfactory management of insect pests when the use of extensive monocultures is continued. In this paper, experiments are suggested that would explore the effectiveness of polycultures or chemotype mixtures with respect to ameliorating the damage of insects on plantation productivity. In addition, mitigating the effects of some insects on plantation productivity by maintaining vigour of fast‐growing eucalypts should be considered.

     

The Nicotiana attenuata LECTIN RECEPTOR KINASE 1 is involved in the perception of insect feeding

The Nicotiana attenuata LECTIN RECEPTOR KINASE 1 (LecRK1) has been recently identified as a component of the mechanism used by plants to suppress the Manduca sexta-triggered accumulation of salicylic acid (SA). The suppression of the SA burst by LecRK1 allows for the unfettered induction of jasmonic acid (JA)-mediated defense responses against M. sexta herbivory. LecRK1 contains a multi-domain extracellular region composed of a G-type Lectin domain and a PAN-AP domain separated by a variable sequence with low similarity to an EGF domain. The LecRK1 intracellular region is composed of a single domain structure with predicted Ser/Thr protein kinase activity. The multi-domain structure of the extracellular region of LecRK1 adds a level of complexity in terms of the potential ligands that this receptor protein could recognize.     

Volatile organic compounds released by maize following herbivory or insect extract application and communication between plants

To protect themselves from herbivory, plants have evolved an arsenal of physical and chemical defences and release a variety of volatile organic compounds (VOCs). By releasing these VOCs, a signalling plant can both reduce herbivory, sometimes by more than 90%, and also warn neighbouring plants about an attack. The aim of this study was to assess the influence of herbivory and insect extract application on VOC release by damaged/treated and nearby undamaged/untreated maize plants. We confirmed that European corn borer (Ostrinia nubilalis) larvae attack or larvae extract application induced maize VOC release. Greater amounts of (Z)‐3‐hexenal, (E)‐2‐hexenal, (Z)‐3‐hexen‐1‐ol, (E)‐2‐hexen‐1‐ol, β‐myrcene, (Z)‐3‐hexen‐1‐yl acetate, 1‐hexyl acetate, (Z)‐ocimene, linalool, benzyl acetate, methyl salicylate, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene, (E)‐β‐farnesene and (Z)‐3‐hexenal, (Z)‐3‐hexen‐1‐ol, (Z)‐3‐hexen‐1‐yl acetate, (Z)‐ocimene, linalool, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene and (E)‐β‐farnesene were released as a result of biotic stress after insect attack or insect extract application. The amounts of each VOC released were qualitatively and quantitatively distinct and dependent on time after biotic stress exposure. However, for all biotic stresses, significantly lower VOC induction was measured when leaves were damaged/treated for three days, as compared to seven days. Our work also demonstrated that undamaged/untreated neighbouring plants also release significant amounts of VOCs. This suggests that VOC emission by a damaged/treated plant stimulates VOC induction in nearby undamaged/untreated plants. However, the concentrations of all VOCs released by neighbouring undamaged/untreated maize plants were lower than those from damaged/treated plants and were negatively correlated with distance from a damaged/treated plant. Still, significant VOC induction occurred in undamaged/untreated plants even at 3 m distance from a damaged/infected plant. Our work suggests that maize plant protective defence responses (VOC emission) can be induced via application of European corn borer extracts.     

Plant water stress and previous herbivore damage affect insect performance

1. Short‐term changes in plant resistance traits can be affected by abiotic factors or damage by herbivores, although how the combined effects of abiotic factors and previous damage affect subsequent insect larval development is not well understood. 2. Complementary glasshouse and field experiments were conducted to evaluate whether plant water stress and previous herbivore damage influenced monarch (Danaus plexippus) larval development on common milkweed, Asclepias syriaca. 3. In the glasshouse, water stress altered a suite of A. syriaca functional traits but did not affect nutrient content, whereas herbivore damage increased leaf nitrogen (N) and reduced the carbon:nitrogen (C:N) ratio. A bioassay experiment showed that monarch larval survival was lower on well‐watered plants that were previously damaged by monarch larva than on damaged and drought‐stressed plants. Bioassay larvae consumed less leaf tissue of previously damaged plants, whereas monarch larval mass was affected additively by water stress and previous damage, after correcting for the amount of leaf tissue consumed. 4. In a 2‐year field experiment, monarch larval performance was higher on previously damaged A. syriaca plants that received experimentally reduced rainfall, relative to plants receiving ambient rainfall. 5. Collectively, these results from glasshouse and field experiments suggest that insect performance was highest on previously damaged plants under water stress and highlight the additive and interactive roles of abiotic and biotic factors on herbivore performance.     

虫害诱导植物间接防御反应的激发与信号转导途径

植物通过产生和释放挥发性物质增加植食性昆虫的天敌对其寄主或猎物的定位,减少植食性昆虫对植物的取食,从而达到间接防御的目的。植物对植食性昆虫所做出间接防御反应激发因子和信号转导途径的研究,对应用虫害诱导植物挥发物引诱害虫天敌,并进一步从植物、植食性昆虫及其天敌间三级营养关系,研究动植物协同进化机理和病虫害防治具有深远意义。本文根据国内外最新研究进展,对虫害诱导植物间接防御反应的激发因子,昆虫取食信号的转导途径及对植物间接防御相关基因的激活等方面进行了系统地综述。     

Adaptations to water gradient in three Rorippa plant species correspond with plant resistance against insect herbivory under drought and waterlogged conditions

1. Plants live in environments where they are constantly, and often simultaneously, exposed to different types of biotic and abiotic stress, such as insect herbivory and water availability. How plants are adapted to abiotic conditions may determine how a surplus or shortage of water affects plant resistance to insect herbivory. Moreover, this effect may vary depending on the feeding mode of the herbivore.
2. We explored how three closely related Rorippa plant species that vary in adaptations to different water levels, resist herbivory by four different insects (aphids: Myzus persicae, Lipaphis erysimi, and caterpillars: Pieris brassicae, Plutella xylostella) under waterlogging or drought conditions. We hypothesized that plants that are differently adapted to water availability will be disparately affected by water availability in their resistance to insect herbivory.
3. On the semi-aquatic plant species Rorippa amphibia, both aphid species reached a larger colony size under drought conditions. This indicates that R. amphibia was compromised in resistance to aphid feeding when under drought conditions, to which it is less well adapted. Water conditions did not affect aphid performance on the flood-plain species Rorippa palustris. On the terrestrial plant species Rorippa sylvestris, aphids performed worse on waterlogged than drought-treated plants. Neither caterpillar species was significantly affected by the water availability of their food plant.
4. Our findings suggest that water availability can have distinct effects on plant–insect interactions. We propose that plant adaptations to water conditions can be a major predictor towards explaining the variation of effects that water availability can have on plant–insect interactions.

     

Behavioural and community ecology of plants that cry for help

Plants respond to insect herbivory with the production of volatiles that attract carnivorous enemies of the herbivores, a phenomenon called indirect defence or 'plants crying for help'. Plants are under selection to maximize Darwinian fitness, and this can be done by making the right 'decisions' (i.e. by responding to environmental stress in ways that maximize seed production). Plant decisions related to the response to herbivory in terms of the emission of herbivore-induced volatiles include 'to respond or not to respond', 'how fast to respond', 'how to respond' and 'when to stop responding'. In this review, the state-of-the-art of the research field is presented in the context of these decisions that plants face. New questions and directions for future research are identified. To understand the consequences of plant responses in a community context, it is important to expand research from individual interactions to multispecies interactions in a community context. To achieve this, detailed information on underlying mechanisms is essential and first steps on this road have been made. This selective review addresses the ecology of herbivore-induced plant volatiles (HIPVs) by integrating information on mechanisms and ecological functions. New questions are identified as well as challenges for extending current information to community ecology.     

植物对虫害的超越补偿作用

阐述了超越补偿作用的本质及其生物学机制,并探讨了超越补偿在生理生态学及栽培学方面的意义,为超越补偿理论在农作物栽培和农、林害虫防治等方面的应用提供科学依据,以进一步提高农、林业生产的经济效益。     

Simultaneous feeding by aboveground and belowground herbivores attenuates plant-mediated attraction of their respective natural enemies

Herbivore-damaged plants emit volatile organic compounds that attract natural enemies of the herbivores. This form of indirect plant defence occurs aboveground as well as belowground, but it remains unclear how simultaneous feeding by different herbivores attacking leaves and roots may affect the production of the respective defence signals. We employed a setup that combines trapping of volatile organic signals and simultaneous measurements of the attractiveness of these signals to above and belowground natural enemies. Young maize plants were infested with either the foliar herbivore Spodoptera littoralis , the root herbivore Diabrotica virgifera virgifera , or with both these important pest insects. The parasitic wasp Cotesia marginiventris and the entomopathogenic nematode Heterorhabditis megidis were strongly attracted if their respective host was feeding on a plant, but this attraction was significantly reduced if both herbivores were on a plant. The emission of the principal root attractant was indeed reduced due to double infestation, but this was not evident for the leaf volatiles. The parasitoid showed an ability to learn the differences in odour emissions and increased its response to the odour of a doubly infested plant after experiencing this odour during an encounter with hosts. This first study to measure effects of belowground herbivory on aboveground tritrophic signalling and vice-versa reemphasizes the important role of plants in bridging interactions between spatially distinct components of the ecosystem.     

The nutritional landscape of host plants for a specialist insect herbivore

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The Lipid A substructure of the Sinorhizobium meliloti lipopolysaccharides is sufficient to suppress the oxidative burst in host plants

Medicago sativa (alfalfa), Medicago truncatula and Nicotiana tabacum cell suspension cultures, responding to elicitation with the production of reactive oxygen species (ROS), were used to analyse the suppressor (and elicitor) activity of lipopolysaccharides (LPS) of the symbiotic soil bacterium Sinorhizobium meliloti. In order to identify the epitopes of the LPS molecule recognized by the plant, S. meliloti mutants defective in LPS biosynthesis and hydrolytically obtained Lipid A were analysed for biological activity. Lipopolysaccharides isolated from Sinorhizobium meliloti mutants 6963 (altered core region) and L994 (no long-chain fatty acid) showed the same ability to suppress the oxidative burst in host plant cell cultures as the wild-type LPS. Lipid A also displayed the same suppressor activity. By contrast, rhizobial LPS, but not Lipid A, was active as an inducer of the oxidative burst reaction in cell cultures of the nonhost Nicotiana tabacum. In host plants of Sinorhizobium meliloti the Lipid A part is sufficient to suppress the oxidative burst, but in non-host plants at least some sugars of the LPS core region are required to induce defence reactions.     

Isopentylamine is a novel defence compound induced by insect feeding in rice

Plants produce a broad variety of defensive metabolites to protect themselves against herbivorous insects. Although polyamines have been implicated in various responses to abiotic and biotic stress, there have been no studies focused on amines in response to insect herbivory. By screening for bioactive amines, we identified isopentylamine as a novel type of herbivory‐induced compound in rice leaves, which was derived from the amino acid leucine in stable isotope labelling experiments. Accumulation of isopentylamine increased during herbivory by the brown planthopper (Nilaparvata lugens, BPH) and the rice‐feeding armyworm (Mythimna loreyi), as well as in response to treatment with the plant hormone, jasmonic acid. Likewise, isopentylamine accumulation was compromised in rice jasmonate biosynthesis mutants, hebiba and Osjar1. In bio‐assays, BPH insects feeding on rice seedlings submerged in 50 mg/L isopentylamine solution had a higher mortality compared with BPH feeding on seedlings submerged in water. Notably, the rice leaves submerged in 50 mg/L solution showed the endogenous concentrations of isopentylamine similar to that induced by BPHs. These results suggest that isopentylamine functions as a new type of plant defence metabolite that is rapidly induced by herbivore attack and deters insect herbivores in rice.     

Indirect plant defense against insect herbivores: a review

Plants respond to herbivore attack by launching 2 types of defenses: direct defense and indirect defense. Direct defense includes all plant traits that increase the resistance of host plants to insect herbivores by affecting the physiology and/or behavior of the attackers. Indirect defense includes all traits that by themselves do not have significant direct impact on the attacking herbivores, but can attract natural enemies of the herbivores and thus reduce plant loss. When plants recognize herbivore‐associated elicitors, they produce and release a blend of volatiles that can attract predators, parasites, and other natural enemies. Known herbivore‐associated elicitors include fatty acid–amino acid conjugates, sulfur‐containing fatty acids, fragments of cell walls, peptides, esters, and enzymes. Identified plant volatiles include terpenes, nitrogenous compounds, and indoles. In addition, constitive traits including extrafloral nectars, food bodies, and domatia can be further induced to higher levels and attract natural enemies as well as provide food and shelter to carnivores. A better understanding of indirect plant defense at global and componential levels via advanced high throughput technologies may lead to utilization of indirect defense in suppression of herbivore damage to plants.     

Studying how plants defend themselves: a chemical weapon produced by chilli fruit

Students often prefer to study animals rather than plants, because they see plants as passive, less interesting organisms. This paper proposes a simple hands-on laboratory exercise for high-school students (grade 12) to arouse their interest in learning about plants and to demonstrate to them that plants are active organisms capable of defending themselves. In this classroom exercise, students investigate how a plant responds to an invader. As a model for study, the exercise uses three organisms: red chilli fruit, the yeast biological control agent Saccharomyces cerevisiae, and the pathogenic fungus Rhizopus stolonifer. Students gain basic knowledge about how plants defend themselves against pathogens by observing the physical changes of an infected wound site and by investigating the chemical compounds produced by plants in response to pathogen invasion; they are also encouraged to think critically about using biological control as a means to promote environmentally friendly agricultural practices.     

Signal transduction downstream of salicylic and jasmonic acid in herbivory-induced parasitoid attraction by Arabidopsis is independent of JAR1 and NPR1

Plants can defend themselves indirectly against herbivores by emitting a volatile blend upon herbivory that attracts the natural enemies of these herbivores, either predators or parasitoids. Although signal transduction in plants from herbivory to induced volatile production depends on jasmonic acid (JA) and salicylic acid (SA), the pathways downstream of JA and SA are unknown. Use of Arabidopsis provides a unique possibility to study signal transduction by use of signalling mutants, which so far has not been exploited in studies on indirect plant defence. In the present study it was demonstrated that jar1‐1 and npr1‐1 mutants are not affected in caterpillar (Pieris rapae)‐induced attraction of the parasitoid Cotesia rubecula. Both JAR1 and NPR1 (also known as NIM1) are involved in signalling downstream of JA in induced defence against pathogens such as induced systemic resistance (ISR). NPR1 is also involved in signalling downstream of SA in defence against pathogens such as systemic acquired resistance (SAR). These results demonstrate that signalling downstream of JA and SA differs between induced indirect defence against herbivores and defence against pathogens such as SAR and ISR. Furthermore, it was demonstrated that herbivore‐derived elicitors are involved in induced attraction of the parasitoid Cotesia rubecula     

Adaptation to host plants may prevent rapid insect responses to climate change

We must consider the role of multitrophic interactions when examining species' responses to climate change. Many plant species, particularly trees, are limited in their ability to shift their geographic ranges quickly under climate change. Consequently, for herbivorous insects, geographic mosaics of host plant specialization could prohibit range shifts and adaptation when insects become separated from suitable host plants. In this study, we examined larval growth and survival of an oak specialist butterfly (Erynnis propertius) on different oaks (Quercus spp.) that occur across its range to determine if individuals can switch host plants if they move into new areas under climate change. Individuals from Oregon and northern California, USA that feed on Q. garryana and Q. kelloggii in the field experienced increased mortality on Q. agrifolia, a southern species with low nutrient content. In contrast, populations from southern California that normally feed on Q. agrifolia performed well on Q. agrifolia and Q. garryana and poorly on the northern, high elevation Q. kelloggii. Therefore, colonization of southern E. propertius in higher elevations and some northern locales may be prohibited under climate change but latitudinal shifts to Q. garryana may be possible. Where shifts are precluded due to maladaptation to hosts, populations may not accrue warm‐adapted genotypes. Our study suggests that, when interacting species experience asynchronous range shifts, historical local adaptation may preclude populations from colonizing new locales under climate change.     

Bidirectional plant‐mediated interactions between rhizobacteria and shoot‐feeding herbivorous insects: a community ecology perspective

1. Plants interact with various organisms, aboveground as well as belowground. Such interactions result in changes in plant traits with consequences for members of the plant‐associated community at different trophic levels. Research thus far focussed on interactions of plants with individual species. However, studying such interactions in a community context is needed to gain a better understanding.
2. Members of the aboveground insect community induce defences that systemically influence plant interactions with herbivorous as well as carnivorous insects. Plant roots are associated with a community of plant‐growth promoting rhizobacteria (PGPR). This PGPR community modulates insect‐induced defences of plants. Thus, PGPR and insects interact indirectly via plant‐mediated interactions.
3. Such plant‐mediated interactions between belowground PGPR and aboveground insects have usually been addressed unidirectionally from belowground to aboveground. Here, we take a bidirectional approach to these cross‐compartment plant‐mediated interactions.
4. Recent studies show that upon aboveground attack by insect herbivores, plants may recruit rhizobacteria that enhance plant defence against the attackers. This rearranging of the PGPR community in the rhizosphere has consequences for members of the aboveground insect community. This review focusses on the bidirectional nature of plant‐mediated interactions between the PGPR and insect communities associated with plants, including (a) effects of beneficial rhizobacteria via modification of plant defence traits on insects and (b) effects of plant defence against insects on the PGPR community in the rhizosphere. We discuss how such knowledge can be used in the development of sustainable crop‐protection strategies.

     

植物源昆虫拒食活性物质的研究和应用

植物在与昆虫协调进化过程中形成了许多具有防御功能的次生代谢物质。作者对近年来植物源昆虫拒食活性物质的研究和应用进行了综述,其结构类型主要涉及萜类、生物碱类、酚类等,讨论了制约植物源昆虫拒食剂发展的因素并展望了植物源昆虫拒食剂在未来害虫综合治理中的作用。