Plant volatiles as method of communication

Plants emit volatile compounds that can act as a communication method to insects, neighboring plants and pathogens. Plants respond to leaf and root damage by herbivores and pathogens by emitting these compounds. The volatile compounds can deter the herbivores or pathogens directly or indirectly by attracting their natural enemies to kill them. The simultaneous damage of plants by herbivores and pathogens can influence plant defense. The induced plant volatiles can also make neighboring plants ready for defense or induce defense in parts distant from the damaged area of the same plant. Belowground root herbivory can alter the defense response to aboveground leaf herbivory. In addition, most plants normally emit volatile compounds from their flowers that directly attract foraging mutualistic insects for nectar, which in turn perform the very important function of pollination for subsequent reproduction. The volatile compounds emitted from the floral and vegetative parts of plants belong to three main classes of compounds: terpenoids, phenylpropanoids/benzenoids, and C6-aldehydes (green-leaf volatiles). The volatile phytohormones methyl salicylate and methyl jasmonate serve as important signaling molecules for communication purposes, and interact with each other to optimize the plant defense response. Here we discuss and integrate the current knowledge on all types of communication between plants and insects, neighboring plants and pathogens that are mediated through plant volatiles.     

丛枝菌丝网络介导苜蓿植株间机械损伤信号的传递

为了解植株间的菌丝网络(common mycorrhizal networks,CMNs)的功能,对丛枝菌根菌丝网络在紫花苜蓿(Medicago sativa)机械损伤信号传递中的作用进行了研究.结果表明,与经机械损伤处理的供体植株有CMNs连接的受体植株叶片中挥发物的数量和种类都显著高于无CMNs连接的植株;供体植株...     

Biosynthèse des glucosinolates indoliques et rôle écologique de leurs modifications secondaires

Indole glucosinolates are plant secondary metabolites derived from the amino acid tryptophan. They are part of a large group of sulfur-containing molecules almost exclusively found among Brassicales, which include the mustard family (Brassicaceae) with many edible plant species of major nutritional importance. These compounds mediate numerous interactions between these plants and their natural enemies and are therefore of major biological and economical interest. This literature review aims at taking stock of recent advances of our knowledge about the biosynthetic pathways of indole glucosinolates, but also about the defense strategies and ecological processes involving these metabolites.     

Differences in volatile terpene composition between the bark and leaves of tropical tree species

Volatile terpenes are among the most diverse class of defensive compounds in plants, and they are implicated in both direct and indirect defense against herbivores. In terpenes, both the quantity and the diversity of compounds appear to increase the efficiency of defense as a diverse blend of compounds provides a more efficient protection against a broader range of herbivores and limits the chances that an enemy evolves resistance. Theory predicts that plant defensive compounds should be allocated differentially among tissues according to the value of the tissue, its cost of construction and the herbivore pressure on it. We collected volatile terpenes from bark and leaves of 178 individual tree belonging to 55 angiosperm species in French Guiana and compare the kind, amount, and diversity of compounds in these tissues. We hypothesized that in woody plants, the outermost part of the trunk should hold a more diverse blend of volatile terpenes. Additionally, as herbivore communities associated with the leaves is different to the one associated with the bark, we also hypothesized that terpene blends should be distinct in the bark vs. the leaves of a given species. We found that the mixture of volatile terpenes released by bark is different and more diverse than that released by leaves, both in monoterpenes and sesquiterpenes. This supports our hypothesis and further suggests that the emission of terpenes by the bark should be more important for trunk defense than previously thought.     

Effects of feeding Spodoptera littoralis on lima bean leaves. II. Continuous mechanical wounding resembling insect feeding is sufficient to elicit herbivory-related volatile emission

Herbivore feeding elicits defense responses in infested plants, including the emission of volatile organic compounds that can serve as indirect defense signals. Until now, the contribution of plant tissue wounding during the feeding process in the elicitation of defense responses has not been clear. For example, in lima bean (Phaseolus lunatus), the composition of the volatiles induced by both the insect caterpillar Spodoptera littoralis and the snail Cepaea hortensis is very similar. Thus, a mechanical caterpillar, MecWorm, has been designed and used in this study, which very closely resembles the herbivore-caused tissue damage in terms of similar physical appearance and long-lasting wounding period on defined leaf areas. This mode of treatment was sufficient to induce the emission of a volatile organic compound blend qualitatively similar to that as known from real herbivore feeding, although there were significant quantitative differences for a number of compounds. Moreover, both the duration and the area that has been mechanically damaged contribute to the induction of the whole volatile response. Based on those two parameters, time and area, which can replace each other to some extent, a damage level can be defined. That damage level exhibits a close linear relationship with the accumulation of fatty acid-derived volatiles and monoterpenes, while other terpenoid volatiles and methyl salicylate respond in a nonlinear manner. The results strongly suggest that the impact of mechanical wounding on the induction of defense responses during herbivore feeding was until now underestimated. Controlled and reproducible mechanical damage that strongly resembles the insect's feeding process represents a valuable tool for analyzing the role of the various signals involved in the induction of plant defense reactions against herbivory.     

The Anomeric Equilibrium of Glucose in Acidic and Basic Media

In order to elucidate the nature of malodor from piggery wastes, volatile compounds in fresh faeces, fresh urine, rotten urine, and rotten mixtures of faeces and urine were isolated by freeze vacuum distillation and continuous extraction and identified by gas chromatography-mass spectrometry. Many alcohols were detected not in fresh urine, but in faeces. Various fatty acids were determined at high concentrations in all samples, but their abundance was different in faeces and urine. Large amounts of phenols came from urine. Aromatic carboxylic acids were detected only in urine and decreased rapidly during digestion. Indole and 3-methylindole which were present only in faeces showed a reverse change of concentration during digestion.     

Metabolic engineering of aroma components in fruits

Plants have the ability to produce a diversity of volatile metabolites, which attract pollinators and seed dispersers and strengthen plant defense responses. Selection by plant breeders of traits such as rapid growth and yield leads, in many cases, to the loss of flavor and aroma quality in crops. How the aroma can be improved without affecting other fruit attributes is a major unsolved issue. Significant advances in metabolic engineering directed at improving the set of volatiles that the fruits emit has been aided by the characterization of enzymes involved in the biosynthesis of flavor and aroma compounds in some fruits. However, before this technology can be successfully applied to modulate the production of volatiles in different crops, further basic research is needed on the mechanisms that lead to the production of these compounds in plants. Here we review the biosynthesis and function of volatile compounds in plants, and the attempts that have been made to manipulate fruit aroma biosynthesis by metabolic engineering. In addition, we discuss the possibilities that molecular breeding offers for aroma enhancement and the implications of the latest advances in biotechnological modification of fruit flavor and aroma.     

Deciphering the Role of Ethylene in Plant–Herbivore Interactions

Most plants emit ethylene in response to herbivory by insects from many different feeding guilds. The elicitors of these ethylene emissions are thought to be microorganisms or oral secretion-specific compounds that are transferred when the attacking insect feeds. To find the receptors for these elicitors and describe the signaling cascades that are subsequently activated will be the challenge of future research. Past experiments on the function of herbivore-induced ethylene, which were biased toward the use of chemical treatments to manipulate ethylene, identified seven ethylene-dependent defense responses. In contrast, a genetic toolbox that consists of several mutants has rarely been used and to date, mutants have helped to identify only one additional ethylene-dependent defense response. Ethylene-dependent responses include the emission of specific volatile organic compounds as indirect defense, the accumulation of phenolic compounds, and proteinase inhibitor activity. Besides being ethylene regulated, these defenses depend strongly on the wound-hormone jasmonic acid (JA). That ethylene requires the concomitant induction of JA, or other signals, appears to be decisive. Rather than being the principal elicitor of defense responses, ethylene modulates the sensitivity to a second signal and its downstream responses. Given this modulator role, and the artifacts associated with the use of chemical treatments to manipulate ethylene production and perception, future advances in the study of ethylene’s function in plant–herbivore interactions will likely come from the use of signaling mutants or transgenic plants. It will be exciting to see if adaptive phenotypic plasticity is largely an ethylene-mediated response.     

Lipoxygenase H1 gene silencing reveals a specific role in supplying fatty acid hydroperoxides for aliphatic aldehyde production.

Lipoxygenases catalyze the formation of fatty acid hydroperoxide precursors of an array of compounds involved in the regulation of plant development and responses to stress. To elucidate the function of the potato 13-lipoxygenase H1 (LOX H1), we have generated transgenic potato plants with reduced expression of the LOX H1 gene as a consequence of co-suppression-mediated gene silencing. Three independent LOX H1-silenced transgenic lines were obtained, having less than 1% of the LOX H1 protein present in wild-type plants. This depletion of LOX H1 has no effect on the basal or wound-induced levels of jasmonates derived from 13-hydroperoxylinolenic acid. However, LOX H1 depletion results in a marked reduction in the production of volatile aliphatic C6 aldehydes. These compounds are involved in plant defense responses, acting as either signaling molecules for wound-induced gene expression or as antimicrobial substances. LOX H1 protein was localized to the chloroplast and the protein, expressed in Escherichia coli, showed activity toward unesterified linoleic and linolenic acids and plastidic phosphatidylglycerol. The results demonstrate that LOX H1 is a specific isoform involved in the generation of volatile defense and signaling compounds through the HPL branch of the octadecanoid pathway.     

吲哚——种间及跨界信号分子新成员

吲哚作为一种典型的氮杂环芳烃化合物,在自然界中广泛存在。近年来,越来越多的研究表明吲哚具有一定的生物活性,是一种新型种间及跨界的信号分子。研究发现,吲哚不仅可以调节微生物的毒性、耐药性、生物膜形成以及群感效应等生理生化行为,调控植物生长发育和防御系统的形成过程,还能够影响动物的肠道炎症、细胞氧化压力及荷尔蒙分泌等生理健康。因此吲哚在微生物代谢、动物健康和植物生长等多个方面扮演了重要角色,具有重要的生物学及生态学双重意义。文中综述了吲哚从生物代谢到信号传递的研究历史,及其在微生物种内或种间以及微生物-动植物之间跨界的信号传导与调控作用的研究进展,旨在为揭示复杂环境中吲哚生物代谢及信号调控的生物学意义与生态学机制提供重要的理论指导。     

IAN-bildung aus glucobrassicin: pH-Abhängigkeit und wachstumsphysiologische Bedeutung

Summary Indole auxin synthesis during enzymatic hydrolysis of glucobrassicin by myrosinase proved to be strictly dependent on pH. Neither IAN nor other indole compounds with auxin activity are synthesized at pH values higher than 5.2. An in vivo function of the indole glucosinolates as auxin precursors in Cruciferae is therefore of low probability.     

Metabolic engineering of plant volatiles

Metabolic engineering of the volatile spectrum offers enormous potential for plant improvement because of the great contribution of volatile secondary metabolites to reproduction, defense and food quality. Recent advances in the identification of the genes and enzymes responsible for the biosynthesis of volatile compounds have made this metabolic engineering highly feasible. Notable successes have been reported in enhancing plant defenses and improving scent and aroma quality of flowers and fruits. These studies have also revealed challenges and limitations which will be likely surmounted as our understanding of plant volatile network improves.     

Physiological function and ecological aspects of fatty acid-amino acid conjugates in insects†

In tritrophic interactions, plants recognize herbivore-produced elicitors and release a blend of volatile compounds (VOCs), which work as chemical cues for parasitoids or predators to locate their hosts. From detection of elicitors to VOC emissions, plants utilize sophisticated systems that resemble the plant–microbe interaction system. Fatty acid–amino acid conjugates (FACs), a class of insect elicitors, resemble compounds synthesized by microbes in nature. Recent evidence suggests that the recognition of insect elicitors by an ancestral microbe-associated defense system may be the origin of tritrophic interactions mediated by FACs. Here we discuss our findings in light of how plants have customized this defense to be effective against insect herbivores, and how some insects have successfully adapted to these defenses.     

Acyltransferases in plants: a good time to be BAHD

Acylation is a common and biochemically significant modification of plant secondary metabolites. Plant BAHD acyltransferases constitute a large family of acyl CoA-utilizing enzymes whose products include small volatile esters, modified anthocyanins, as well as constitutive defense compounds and phytoalexins. The catalytic versatility of BAHD enzymes makes it very difficult to make functional predictions from primary sequence alone. Recent advances in genome sequencing and the availability of the first crystal structure of a BAHD member are, however, providing insights into the evolution and function of these acyltransferases within the plant kingdom.     

Systemin and jasmonic acid regulate constitutive and herbivore-induced systemic volatile emissions in tomato, Solanum lycopersicum

Transgenic tomato (Solanum lycopersicum) plants that overexpress the Prosystemin gene (35S::PS) and plants with a mutation in the JA biosynthetic pathway (def1) are known to exhibit a constitutive or reduced wound response, respectively. Here it is demonstrated that several independent 35S::PS lines emit high levels of specific volatiles in addition to increased accumulation of proteinase inhibitors (PIs). Furthermore, the temporal dynamics of systemically induced volatile compounds including green-leaf volatiles, terpenes, and shikimic acid-derivatives from 35S::PS and def1 plants in response to herbivore wounding and treatment with jasmonic acid (JA) are described. Application of JA induced defense protein accumulation and volatile emissions in wild type plants, but did not further increase systemic volatile emissions from 35S::PS plants. Wounding by Manduca sexta larvae induced synthesis of defense proteins and emission of volatiles in wild type plants, but not in def1 plants. Application of jasmonic acid restored the local and systemic accumulation of defense proteins in def1, as well as enhanced herbivore-induced volatile emissions. These results provide strong support for the role of prosystemin- and JA-signaling in the regulation of volatile emissions in tomato plants.     

Volatiles from four Astragalus species: phenological changes and their chemotaxonomical application

This paper shows the changes of the volatile compounds from four Astragalus species at three phenological stages: leaf development, flowering and fructification, which might be connected with the plant defense. After GC/MS analyses of Astragalus glycyphyllos L., A. hamosus L., A. cicer L. and A. spruneri Boiss., different groups of volatile compounds were found: hydrocarbons, alcohols, aldehydes and ketones, esters, terpenes, chlorinated compounds, etc. Identified volatiles were used for a cluster analysis in order to make chemotaxonomic conclusions for these evolutionary different species.     

Response of Pyropia haitanensis to agaro-oligosaccharides evidenced mainly by the activation of the eicosanoid pathway

The marine red alga Pyropia haitanensis (Protoflorideophyceae, Bangiaceae) has a nonvascular and multicellular structure and emerged earlier in evolution than other cultivatable red algae. It has been reported that lipid mediators from both the eicosanoid and octadecanoid pathways are involved in the innate immunity of other marine algae. But the defense strategies of P. haitanensis are not clearly understood. Here, we investigated the lipid defense of P. haitanensis elicited by agaro-oligosaccharides. The results indicate that the resistance of P. haitanensis was elicited and hydrogen peroxide was released by agaro-oligosaccharides. In P. haitanensis, C20 fatty acids are the essential fatty acids. Phospholipase A₂ was activated, and the free fatty acids decreased 3 h after treatment with agaro-oligosaccharides. Gas chromatography–mass spectrometry analyses revealed that the contents of volatile organic compounds increased after treatment for 3 h, which indicated that these free fatty acids were metabolized to volatile organic compounds. In conclusion, the lipid metabolic defense pathway of P. haitanensis was mainly via the C20 metabolism pathway. The C20 fatty acid was rapidly metabolized to volatile organic compounds, but not oxidized to oxylipins in response to agaro-oligosaccharides.     

New insight into the biosynthesis and regulation of indole compounds in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

In spite of their silent and sessile life, plants are dynamic organisms that have developed advanced defence strategies in their adaptation to the pressure of herbivores and pathogens. Natural plant products play an important role as chemical weapons in this warfare. Characteristic of cruciferous plants is the synthesis of nitrogen- and sulphur-rich compounds, such as glucosinolates (Mikkelsen et al. 2002) and indole alkaloids (Pedras et al. 2000). Glucosinolates are believed to be largely non-toxic, but upon tissue disruption, they are hydrolyzed by endogenous -thioglucosidases (myrosinases) (Rask et al. 2000) to primarily isothiocyanates and nitriles, which have many biological activities. These include not only important roles as repellents against herbivorous insects and microorganisms, but also as volatile attraction of specialized insects (Wittstock and Halkier 2002). For humans, these compounds serve as cancer-preventive agents, biopesticides, and flavor compounds (Talalay and Fahey 2001). Indole alkaloids are phytoalexins and production of specific alkaloids is usually limited to only a few species. Cruciferous plants include the model plant Arabidopsis, which produces the indole alkaloid camalexin. This review will focus on the central role of indole-3-acetaldoxime (IAOx) in the biosynthesis of indole glucosinolates, camalexin, and the phytohormone IAA.