Synthetic Herbivore-induced Plant Volatiles Increase Field Captures of Parasitic Wasps

Field evidence for attraction of parasitic wasps from the families Encyrtidae and Mymaridae to grapevines baited with synthetic versions of three herbivore-induced plant volatiles (HIPV) is presented. In a replicated experiment conducted in a juice grape vineyard, sticky cards in blocks baited with controlled-release dispensers of methyl salicylate (MeSA), methyl jasmonate (MeJA) or (Z)-3-hexenyl acetate (HA), trapped significantly greater numbers of Metaphycus sp. (Encyrtidae) than cards in unbaited blocks during May–September. Significantly greater numbers of Anagrus spp. (Mymaridae) were trapped in MeSA and MeJA-baited blocks than in unbaited blocks, during August–September. Greater numbers were recorded in HA-baited blocks in September only. Previous studies showed Encyrtidae and Anagrus spp. were not attracted to sticky cards baited with vials of MeSA, MeJA or HA. Possible reasons for attraction in this study are discussed including the possibility that synthetic, gaseous HIPV from controlled-release dispensers may stimulate plants to produce natural blends of parasitoid-attracting volatiles.     

虫害诱导植物挥发物（HIPVs）：从诱导到生态功能

植物被植食性昆虫危害后,其地上和地下部分均能释放虫害诱导植物挥发物(HIPVs).HIPVs不仅能招引植食性昆虫的天敌,而且还可对植食性昆虫和临近植物产生影响,从而调节植物、植食性昆虫、天敌三者之间的相互关系.根据最近的研究结果,主要就HIPVs的诱导物类型、虫害后植物体内的早期信号和信号传导、HIPVs的合成、释放、组成,以及其生态功能等进行了系统性的综述,并提出了今后该领域的研究方向.     

Plant Volatiles: Recent Advances and Future Perspectives

Volatile compounds act as a language that plants use for their communication and interaction with the surrounding environment. To date, a total of 1700 volatile compounds have been isolated from more than 90 plant families. These volatiles, released from leaves, flowers, and fruits into the atmosphere and from roots into the soil, defend plants against herbivores and pathogens or provide a reproductive advantage by attracting pollinators and seed dispersers. Plant volatiles constitute about 1% of plant secondary metabolites and are mainly represented by terpenoids, phenylpropanoids/benzenoids, fatty acid derivatives, and amino acid derivatives. In this review we focus on the functions of plant volatiles, their biosynthesis and regulation, and the metabolic engineering of the volatile spectrum, which results in plant defense improvement and changes of scent and aroma properties of flowers and fruits.     

Herbivore-specific plant volatiles prime neighboring plants for nonspecific defense responses

Plants produce species-specific herbivore-induced plant volatiles (HIPVs) after damage. We tested the hypothesis that herbivore-specific HIPVs prime neighboring plants to induce defenses specific to the priming herbivore. Since Manduca sexta (specialist) and Heliothis virescens (generalist) herbivory induced unique HIPV profiles in Nicotiana benthamiana, we used these HIPVs to prime receiver plants for defense responses to simulated herbivory (mechanical wounding and herbivore regurgitant application). Jasmonic acid (JA) accumulations and emitted volatile profiles were monitored as representative defense responses since JA is the major plant hormone involved in wound and defense signaling and HIPVs have been implicated as signals in tritrophic interactions. Herbivore species-specific HIPVs primed neighboring plants, which produced 2 to 4 times more volatiles and JA after simulated herbivory when compared to similarly treated constitutive volatile-exposed plants. However, HIPV-exposed plants accumulated similar amounts of volatiles and JA independent of the combination of priming or challenging herbivore. Furthermore, volatile profiles emitted by primed plants depended only on the challenging herbivore species but not on the species-specific HIPV profile of damaged emitter plants. This suggests that feeding by either herbivore species primed neighboring plants for increased HIPV emissions specific to the subsequently attacking herbivore and is probably controlled by JA.     

Evolutionary changes in an invasive plant support the defensive role of plant volatiles

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Within-Plant Migration of the Predatory Mite Typhlodromalus aripo from the Apex to the Leaves of Cassava: Response to Day–Night Cycle, Prey Location and Prey Density

Under attack by herbivores, plants produce a blend of “herbivore-induced plant volatiles (HIPV)” that help natural enemies of herbivores locating their prey, thereby helping plants to reduce damage from herbivory. The amount of HIPV emitted by plants increases with herbivore density and is positively correlated with the intensity of the olfactory response of natural enemies. In this study, we determined the effects of density or within-plant distribution of the herbivorous mite Mononychellus tanajoa on movement of the predatory mite Typhlodromalus aripo out of apices of cassava plants. Proportions of T. aripo that migrated out of apex, and distances traveled were significantly higher when M. tanajoa was further away from the apex—i.e. on middle or bottom leaves of cassava plants—than when present on top leaves, or absent from the plant. This supports previous field observations that T. aripo is not a sit-and-wait predator but uses HIPV to search and locate its prey within cassava plant.     

Herbivore-induced responses and patch heterogeneity affect abundance of arthropods on plants

Abstract.  1. Plants respond to herbivore damage by inducing defences that can affect the abundance of herbivores and predators. These tritrophic interactions may be influenced by heterogeneity in plant neighbourhood.
2. In the present study, the effects of induced responses on the abundance of herbivores (flea beetles and aphids), omnivores (pirate bugs and thrips), and predators (lady beetles and spiders) on individual plants and their neighbours between and within patches composed of three tomato plants was investigated.
3. Herbivore damage was manipulated to create homogeneous patches where either all or none of the plants had defences induced by herbivore damage, and heterogeneous patches where only one of the plants was induced.
4. Arthropod abundance on plants at different scales was compared by testing between patch effects (patch level), for neighbourhood effects at the plant phenotype level (neighbourhood level), and between near and far plants (within patch position).
5. At the patch level , plants in homogeneously induced patches contained fewer flea beetles and pirate bugs, but more lady beetles, compared with homogeneously non-induced patches. There was no effect of patch type on the abundance of aphids, thrips, and spiders on plants.
6. At the neighbourhood level , induced plants in heterogeneous patches contained more flea beetles and pirate bugs compared with induced plants in homogeneous patches, indicating that the abundance of some herbivores and omnivores on induced plants varied depending on the phenotype of the other plants within the patch. Within patch position, there was no evidence that the abundance of herbivores or predators on non-induced plants was affected by proximity to an induced plant.
7. Therefore, variation in plant neighbourhood generated by induced plant responses affected the abundance of three arthropods from three feeding guilds.     

植物挥发性物质及其代谢工程

植物挥发性物质在植物之间和植物与昆虫间的化学通讯中起着重要作用.有关这些次生物质的生物合成、代谢调控、生理功能以及与环境相互作用的研究近十多年来取得了重要进展.迄今为止,已经有3 0多种植物挥发性物质的合成酶基因被克隆.这些基因调控着植物萜类、芳香化合物、脂肪酸衍生物这三大类主要挥发性物质的生物合成.由于潜在的应用价值,近几年该领域颇受注目,特别是应用基因工程技术设计植物释放特殊气味物质,诸如特定的驱避剂或者其它控制植物或昆虫行为的特殊气味乃至与人类健康相关的药用气味物质.该文就植物挥发性物质的生物合成、生理和生态功能以及基因工程方面的研究进展作一概述.     

虫害诱导挥发物的生态调控功能

虫害诱导挥发物（herbivore-induced plant volatiles, HIPVs）是植物受害虫胁迫后释放的挥发性物质,是植物与周围环境进行信息交流的媒介。环境中的天敌、害虫和植物通过感知HIPVs所携带的信息,对各自的行为或生理生化反应做出相应的调整。介绍了挥发物的种类及主要的生物合成途径,概括了影响天敌依据HIPVs搜寻寄主和猎物的主要因素。综述了这类挥发性物质对植食性昆虫寄主选择或产卵行为的影响,介绍了植物地上部分和地下部分受害后对彼此间接防御的影响,讨论了多种害虫加害同种植物后对天敌搜寻猎物或寄主行为的影响。另外,作为损伤信号,HIPVs还能诱导同株植物未受害部位和邻近植株的防御反应。最后,对HIPVs在害虫防治中的应用现状及前景作了介绍和讨论。     

稻飞虱卵期寄生蜂稻虱缨小蜂引诱剂的筛选与田间试验(英文)

植食性昆虫的天敌能够利用虫害诱导的挥发物进行有效的寄主或猎物定位。为了开发稻飞虱卵期天敌稻虱缨小蜂Anagrus nilaparvatae Pang et Wang的引诱剂,分别在室内和室外检测了多种褐飞虱诱导的水稻挥发物组分对褐飞虱卵期天敌稻虱缨小蜂的引诱作用。Y型嗅觉仪实验结果表明,5种单一化合物,Z-3-己烯乙酸酯,1-戊烯基-3-醇,Z-3-己烯醛,芳樟醇和水杨酸甲酯,以及3种混合物,水杨酸甲酯+Z-3-己烯醛,Z-3-己烯醛+Z-3-己烯乙酸酯+芳樟醇,水杨酸甲酯+Z-3-己烯醛+Z-3-己烯乙酸酯+芳樟醇,对稻虱缨小蜂具有明显引诱作用。田间试验表明,3种单一化合物,Z-3-己烯乙酸酯,Z-3-己烯醛和芳樟醇,以及一种混合物,水杨酸甲酯+Z-3-己烯醛+Z-3-己烯乙酸酯+芳樟醇,能明显提高稻虱缨小蜂对褐飞虱卵的寄生作用。这些结果对于改善褐飞虱治理具有重要的意义。     

Visualization of Vascular Ca2+ Signaling Triggered by Paracrine Derived ROS

Oxidative stress has been implicated in a number of pathologic conditions including ischemia/reperfusion damage and sepsis. The concept of oxidative stress refers to the aberrant formation of ROS (reactive oxygen species), which include O₂^•-, H₂O₂, and hydroxyl radicals. Reactive oxygen species influences a multitude of cellular processes including signal transduction, cell proliferation and cell death^1-6. ROS have the potential to damage vascular and organ cells directly, and can initiate secondary chemical reactions and genetic alterations that ultimately result in an amplification of the initial ROS-mediated tissue damage. A key component of the amplification cascade that exacerbates irreversible tissue damage is the recruitment and activation of circulating inflammatory cells. During inflammation, inflammatory cells produce cytokines such as tumor necrosis factor-α (TNFα) and IL-1 that activate endothelial cells (EC) and epithelial cells and further augment the inflammatory response⁷. Vascular endothelial dysfunction is an established feature of acute inflammation. Macrophages contribute to endothelial dysfunction during inflammation by mechanisms that remain unclear. Activation of macrophages results in the extracellular release of O₂^•- and various pro-inflammatory cytokines, which triggers pathologic signaling in adjacent cells⁸. NADPH oxidases are the major and primary source of ROS in most of the cell types. Recently, it is shown by us and others^9,10 that ROS produced by NADPH oxidases induce the mitochondrial ROS production during many pathophysiological conditions. Hence measuring the mitochondrial ROS production is equally important in addition to measuring cytosolic ROS. Macrophages produce ROS by the flavoprotein enzyme NADPH oxidase which plays a primary role in inflammation. Once activated, phagocytic NADPH oxidase produces copious amounts of O₂^•- that are important in the host defense mechanism^11,12. Although paracrine-derived O₂^•- plays an important role in the pathogenesis of vascular diseases, visualization of paracrine ROS-induced intracellular signaling including Ca²⁺ mobilization is still hypothesis. We have developed a model in which activated macrophages are used as a source of O₂^•- to transduce a signal to adjacent endothelial cells. Using this model we demonstrate that macrophage-derived O₂^•- lead to calcium signaling in adjacent endothelial cells.     

虫害诱导的植物挥发物:基本特性、生态学功能及释放机制

植物在遭受植食性昆虫攻击时,能通过释放挥发物调节植物、植食性昆虫及其天敌三者间的相互关系,并由此而防御植食性昆虫。主要就虫害诱导的植物挥发物的基本特性、生态学功能及其释放机制进行了系统性综述,并提出了今后的研究方向。     

Assessing Stomatal Response to Live Bacterial Cells using Whole Leaf Imaging

Stomata are natural openings in the plant epidermis responsible for gas exchange between plant interior and environment. They are formed by a pair of guard cells, which are able to close the stomatal pore in response to a number of external factors including light intensity, carbon dioxide concentration, and relative humidity (RH). The stomatal pore is also the main route for pathogen entry into leaves, a crucial step for disease development. Recent studies have unveiled that closure of the pore is effective in minimizing bacterial disease development in Arabidopsis plants; an integral part of plant innate immunity. Previously, we have used epidermal peels to assess stomatal response to live bacteria (Melotto et al. 2006); however maintaining favorable environmental conditions for both plant epidermal peels and bacterial cells has been challenging. Leaf epidermis can be kept alive and healthy with MES buffer (10 mM KCl, 25 mM MES-KOH, pH 6.15) for electrophysiological experiments of guard cells. However, this buffer is not appropriate for obtaining bacterial suspension. On the other hand, bacterial cells can be kept alive in water which is not proper to maintain epidermal peels for long period of times. When an epidermal peel floats on water, the cells in the peel that are exposed to air dry within 4 hours limiting the timing to conduct the experiment. An ideal method for assessing the effect of a particular stimulus on guard cells should present minimal interference to stomatal physiology and to the natural environment of the plant as much as possible. We, therefore, developed a new method to assess stomatal response to live bacteria in which leaf wounding and manipulation is greatly minimized aiming to provide an easily reproducible and reliable stomatal assay. The protocol is based on staining of intact leaf with propidium iodide (PI), incubation of staining leaf with bacterial suspension, and observation of leaves under laser scanning confocal microscope. Finally, this method allows for the observation of the same live leaf sample over extended periods of time using conditions that closely mimic the natural conditions under which plants are attacked by pathogens.     

Within-plant signalling via volatiles overcomes vascular constraints on systemic signalling and primes responses against herbivores

Plant volatiles play important roles in signalling between plants and insects, but their role in communication among plants remains controversial. Previous research on plant–plant communication has focused on interactions between neighbouring plants, largely overlooking the possibility that volatiles function as signals within plants. Here, we show that volatiles released by herbivore-wounded leaves of hybrid poplar ( Populus deltoides  ×  nigra ) prime defences in adjacent leaves with little or no vascular connection to the wounded leaves. Undamaged leaves exposed to volatiles from wounded leaves on the same stem had elevated defensive responses to feeding by gypsy moth larvae ( Lymantria dispar L.) compared with leaves that did not receive volatiles. Volatile signals may facilitate systemic responses to localized herbivory even when the transmission of internal signals is constrained by vascular connectivity. Self-signalling via volatiles is consistent with the short distances over which plant response to airborne cues has been observed to occur and has apparent benefits for emitting plants, suggesting that within-plant signalling may have equal or greater ecological significance than signalling between plants.     

植物抗虫“防御警备”: 概念、机理与应用

植物抗虫“防御警备”是指受到某些生物或者非生物因子刺激警备后,植物会提前做好抗虫防御准备,之后当再次受到害虫袭击时,植物会产生更加快速和强烈的抗虫防御反应,从而使自身抗虫性显著提高.这是近年来新发现的植物防御害虫的一种策略,是一种特殊的诱导抗虫机制.植食性昆虫的取食、分泌物、产卵、为害诱导的植物挥发物(HIPVs)以及某些有益微生物、植物营养元素、重金属和一些化学物质均可以引起植物产生抗虫防御警备.防御警备具有抗性高效、持久、环境友好,甚至可以遗传到子代等优点.本文综述了近年来有关植物抗虫防御警备的研究,主要概括了植物抗虫防御警备的一般特征、刺激警备因子和形成机制,并对其在生产实践中的应用前景进行了简要分析,提出了这一领域尚未解决的问题和亟待深入的研究方向.通过合适的方法使植物产生抗虫防御警备可以大大减少杀虫剂的使用,成为害虫综合防治的重要手段.     

Host plant defense signaling in response to a coevolved herbivore combats introduced herbivore attack

Defense‐free space resulting from coevolutionarily naïve host plants recently has been implicated as a factor facilitating invasion success of some insect species. Host plants, however, may not be entirely defenseless against novel herbivore threats. Volatile chemical‐mediated defense signaling, which allows plants to mount specific, rapid, and intense responses, may play a role in systems experiencing novel threats. Here we investigate defense responses of host plants to a native and exotic herbivore and show that (1) host plants defend more effectively against the coevolved herbivore, (2) plants can be induced to defend against a newly‐associated herbivore when in proximity to plants actively defending against the coevolved species, and (3) these defenses affect larval performance. These findings highlight the importance of coevolved herbivore‐specific defenses and suggest that naïveté or defense limitations can be overcome via defense signaling. Determining how these findings apply across various host–herbivore systems is critical to understand mechanisms of successful herbivore invasion.     

The chemistry of defense and apparency in the corollas ofNicotiana attenuata

The morphological and chemical characteristics of flowers which attract pollinators present a dilemma for plants; advertising may increase the apparency of plants to their predators and some pollinators are also predators. We explore how a self-compatible disturbance species,Nicotiana attenuata, copes with this potential dilemma by examining the changes in emission of chemicals from flowers in response to pollination and herbivory. We propose that chemical changes induced by herbivory and pollination reflect the function of the chemicals in the plant. The emission of a single compound, benzyl acetone (BA, 4-phenyl-2-butanone), by flowers increases dramatically (50x) in the evening, peaking just after dark —a pattern of emission characteristic of moth-pollinated flowers. Pools of BA were found only in the outer lip of the corolla where pollinators come in contact with the flower, and diurnal changes in the size of the corolla pool closely paralleled the amount emitted by intact flowers throughout the day, as determined by headspace sampling. Pollination dramatically decreases both the pools of BA in the corolla and its emission from flowers. Similarly, nicotine, a broadly biocidal defense metabolite and an induced defense in vegetative and reproductive tissues, is also found in the headspace of flowers and is principally localized in the basal parts of the corolla below the attachment of the filaments and the nectar reward. Moreover, the dynamics of the corolla pools of BA and nicotine throughout the day are consistent with their roles in advertisement and defense, respectively. The corolla pools of nicotine are stable throughout the day except during the period of peak BA production and emission when nicotine pools decrease significantly. The coordinated increase in BA emission and decline in nicotine pools are not inexorably linked, because herbivory or mechanical damage to corolla tissue rapidly increases corolla nicotine pools without affecting the increase in BA pools. Similarly, leaf damage results in a slower, systemic increase in corolla nicotine pools during reproductive growth but again does not affect BA pools. Excised flowers emitted BA in a manner similar to that of intact flowers, and excision of a majority of flowers from a plant did not alter the BA emission patterns of the remaining flowers. We conclude that althoughN. attenuata's defensive and advertisement chemistries respond synchronously to some environmental stimuli, the flowers' chemical responses to pollinators and herbivoresare distinct and the differences reflect their ecological roles. We propose that the cost-benefit framework of the optimal defense and apparency theories can be fruitfully applied to the allocation of defense metabolites and floral volatiles that function in pollinator attraction, and that this framework can be readily tested by manipulating the patterns of the emissions of plants in the field.     

Characterizing Herbivore Resistance Mechanisms: Spittlebugs on Brachiaria spp. as an Example

Plants can resist herbivore damage through three broad mechanisms: antixenosis, antibiosis and tolerance¹. Antixenosis is the degree to which the plant is avoided when the herbivore is able to select other plants². Antibiosis is the degree to which the plant affects the fitness of the herbivore feeding on it¹.Tolerance is the degree to which the plant can withstand or repair damage caused by the herbivore, without compromising the herbivore''s growth and reproduction¹. The durability of herbivore resistance in an agricultural setting depends to a great extent on the resistance mechanism favored during crop breeding efforts³.We demonstrate a no-choice experiment designed to estimate the relative contributions of antibiosis and tolerance to spittlebug resistance in Brachiaria spp. Several species of African grasses of the genus Brachiaria are valuable forage and pasture plants in the Neotropics, but they can be severely challenged by several native species of spittlebugs (Hemiptera: Cercopidae)⁴.To assess their resistance to spittlebugs, plants are vegetatively-propagated by stem cuttings and allowed to grow for approximately one month, allowing the growth of superficial roots on which spittlebugs can feed. At that point, each test plant is individually challenged with six spittlebug eggs near hatching. Infestations are allowed to progress for one month before evaluating plant damage and insect survival. Scoring plant damage provides an estimate of tolerance while scoring insect survival provides an estimate of antibiosis. This protocol has facilitated our plant breeding objective to enhance spittlebug resistance in commercial brachiariagrases⁵.