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1.
The attraction of natural enemies of herbivores by volatile organic compounds as an induced indirect defence has been studied in several plant systems. The evidence for their defensive function originates mainly from laboratory studies with trained parasitoids and predators; the defensive function of these emissions for plants in natural settings has been rarely demonstrated. In native populations and laboratory Y-tube choice experiments with transgenic Nicotiana attenuata plants unable to release particular volatiles, we demonstrate that predatory bugs use terpenoids and green leaf volatiles (GLVs) to locate their prey on herbivore-attacked plants. By attracting predators with volatile signals, this native plant reduces its herbivore load – demonstrating the defensive function of herbivore-induced volatile emissions. However, plants producing GLVs are also damaged more by flea beetles. The implications of these conflicting ecological effects for the evolution of induced volatile emissions and for the development of sustainable agricultural practices are discussed.  相似文献   

2.
Dimethylallyl diphosphate (DMADP) and geranyl diphosphate (GDP) are the last precursors of isoprene and monoterpenes emitted by leaves, respectively. DMADP and GDP pools were measured in leaves of plants emitting isoprene (Populus alba), monoterpenes (Quercus ilex and Mentha piperita), or nonemitting isoprenoids (Prunus persica). Detectable pools were found in all plant species, but P. persica showed the lowest pool size, which indicates a limitation of the whole pathway leading to isoprenoid biosynthesis in nonemitting species. The pools of DMADP and GDP of nonemitting, isoprene-emitting, and monoterpene-emitting species were partially labeled (generally 40%-60% of total carbon-incorporated (13)C) within the same time by which volatile isoprenoids are fully labeled (15 min). This indicates the coexistence of two pools for both precursors, the rapidly labeled pool presumably occurring in chloroplasts and thereby synthesized by the methylerythritol phosphate pathway and the nonlabeled pool presumably located in the cytosol and synthesized by the mevalonic pathway. In M. piperita storing monoterpenes in specialized leaf structures, the GDP pool remained totally unlabeled, indicating either that monoterpenes are totally formed by the mevalonic pathway or that labeling occurs slowly in comparison to the large pool of stored monoterpenes in this plant. The pools of DMADP and GDP increased during the season (from May to July) but decreased when the leaf was darkened or exposed to very high temperature. In the dark, the pool of DMADP of the isoprene-emitting species decreased faster than the pool of GDP. However, after 6 h of darkness, both pools were depleted to about 10% of the pool size in illuminated leaves. This indicates that both the chloroplastic and the cytosolic pools of precursors are depleted in the dark. When comparing measurements over the season and at different temperatures, an inverse correlation was observed between isoprene emission by P. alba and the DMADP pool size and between monoterpene emission by Q. ilex and the GDP pool size. This suggests that the pool size does not limit the emission of isoprenoids. Rather, it indicates that the flux of volatile isoprenoids effectively controls the size of their pools of precursors.  相似文献   

3.
The carbon of the four main monoterpenes emitted by Quercus ilex L. leaves was completely labeled with 13C after a 20-min feeding with 99% 13CO2. This labeling time course is comparable with the labeling time course of isoprene, the terpenoid emitted by other Quercus species and synthesized in leaf chloroplasts. It is also comparable with that of phosphoglyceric acid. Our experiment therefore provides evidence that monoterpenes emitted by Q. ilex are formed from photosynthesis intermediates and may share the same synthetic pathway with isoprene. By analyzing the rate and the distribution of labeling in the different fragments, we looked for evidence of differential carbon labeling in the [alpha]-pinene emitted. However, the labeling pattern was quite uniform in the different fragments, suggesting that the carbon skeleton of the emitted monoterpenes comes from a unique carbon source.  相似文献   

4.
Herbivore-induced plant volatiles (HIPVs) are commonly emitted from plants after herbivore attack1,2. These HIPVs are mainly regulated by the defensive plant hormone jasmonic acid (JA) and its volatile derivative methyl jasmonate (MeJA)3,4,5. Over the past 3 decades researchers have documented that HIPVs can repel or attract herbivores, attract the natural enemies of herbivores, and in some cases they can induce or prime plant defenses prior to herbivore attack. In a recent paper6, I reported that feeding by gypsy moth caterpillars, exogenous MeJA application, and mechanical damage induce the emissions of volatiles from blueberry plants, albeit differently. In addition, blueberry branches respond to HIPVs emitted from neighboring branches of the same plant by increasing the levels of JA and resistance to herbivores (i.e., direct plant defenses), and by priming volatile emissions (i.e., indirect plant defenses). Similar findings have been reported recently for sagebrush7, poplar8, and lima beans9..Here, I describe a push-pull method for collecting blueberry volatiles induced by herbivore (gypsy moth) feeding, exogenous MeJA application, and mechanical damage. The volatile collection unit consists of a 4 L volatile collection chamber, a 2-piece guillotine, an air delivery system that purifies incoming air, and a vacuum system connected to a trap filled with Super-Q adsorbent to collect volatiles5,6,10. Volatiles collected in Super-Q traps are eluted with dichloromethane and then separated and quantified using Gas Chromatography (GC). This volatile collection method was used n my study6 to investigate the volatile response of undamaged branches to exposure to volatiles from herbivore-damaged branches within blueberry plants. These methods are described here. Briefly, undamaged blueberry branches are exposed to HIPVs from neighboring branches within the same plant. Using the same techniques described above, volatiles emitted from branches after exposure to HIPVs are collected and analyzed.  相似文献   

5.
Changes in emission of volatile organic compounds (VOCs) from tomato induced by the fungus Botrytis cinerea were studied in plants inoculated by spraying with suspensions containing B. cinerea spores. VOC emissions were analysed using on-line gas chromatography–mass spectrometry, with a time resolution of about 1 h, for up to 2 days after spraying. Four phases were delimited according to the starting point and the applied day/night rhythm of the experiments. These phases were used to demonstrate changes in VOC flux caused by B. cinerea infestation. Tomato plants inoculated with B. cinerea emitted a different number and amount of VOCs after inoculation compared to control plants that had been sprayed with a suspension without B. cinerea spores. The changes in emissions were dependent on time after inoculation as well as on the severity of infection. The predominant VOCs emitted after inoculation were volatile products from the lipoxygenase pathway (LOX products). The increased emission of LOX products proved to be a strong indicator of a stress response, indicating that VOC emissions can be used to detect plant stress at an early stage. Besides emission of LOX products, there were also increases in monoterpene emissions. However, neither increased emission of LOX products nor of monoterpenes is specific for B. cinerea attack. The emission of LOX products is also induced by other stresses, and increased emission of monoterpenes seems to be the result of mechanical damage induced by secondary stress impacts on leaves.  相似文献   

6.
7.
Maize plants respond to feeding by arthropod herbivores by producing a number of secondary plant compounds, including volatile organic compounds (VOCs). These herbivore-induced VOCs are not only known to attract natural enemies of the herbivores, but they may also prime inducible defences in neighbouring plants, resulting in stronger and faster defence responses in these VOC-exposed plants. Among the compounds that cause this priming effect, green leaf volatiles (GLVs) have received particular attention, as they are ubiquitous and rapidly emitted upon damage. In this study, we investigated their effects under realistic conditions by applying specially devised dispensers to release four synthetic GLVs at physiologically relevant concentrations in a series of experiments in maize fields. We compared the VOC emission of GLV-exposed maize plants to non-exposed plants and monitored the attraction of herbivores and predators, as well as parasitism of the caterpillar Spodoptera frugiperda, the most common herbivore in the experimental maize fields. We found that maize plants that were exposed to GLVs emitted increased quantities of sesquiterpenes compared to non-exposed plants. In several replicates, herbivorous insects, such as adult Diabrotica beetles and S. frugiperda larvae, were observed more frequently in GLV-treated plots and caused more damage to GLV-exposed plants than to non-exposed plants. Parasitism of S. frugiperda was only weakly affected by GLVs and overall parasitism rates of S. frugiperda were similar in GLV-exposed and non-exposed plots. The effects on insect presence depended on the distance from the GLV-dispensers at which the plants were located. The results are discussed in the context of strategies to improve biological control by enhancing plant-mediated attraction of natural enemies.  相似文献   

8.
We studied the effects of water stress, fertilization and time course on foliar volatile terpene emission rates by Quercus ilex and Pinus halepensis in a garden experiment. The terpenes mostly emitted by both species were α -pinene, β -pinene, β -myrcene and Δ3-carene. P. halepensis emission rates (average 31.45 μg g−1 DM h−1) were similar to those of Q. ilex (average 31.71 μg g−1 DM h−1). The effects of drought (reduction to one-third of full watering) and fertilization (250 kg N ha−1, 250 kg P ha−1, or both) were different depending on the species: the drought treatment significantly increased the terpene emissions from Q. ilex by 33%, and the fertilization treatments reduced the terpene emissions from P. halepensis by 38%. Terpene emission rates increased with time course in parallel to raising summer temperatures in P. halepensis and Q. ilex , whose emission rates were temperature related (r = 0.42 and r = 0.68, respectively) and light related (r = 0.32 and r = 0.57, respectively). There was a positive relationship for P. halepensis , and a negative relationship for Q. ilex, between emission rates and relative water contents. No relationship was found between emission rates and N or P foliar concentrations. The results of this study show complex species-specific responses with stronger and faster short-term responses in terpene-non-storing than in storing species and indicate that terpene emissions may significantly change in the warmer, drier and more fertilized conditions predicted for the next decades in the Mediterranean region.  相似文献   

9.
Feeding by the large pine weevil (Hylobius abietis L.) causes severe damage to the bark of Scots pine (Pinus sylvestris L.) seedlings. We measured photosynthesis, the emission of volatile organic compounds from intact and weevil-damaged bark and systemic emissions from undamaged foliage. Feeding damage did not affect photosynthesis. Monoterpenes dominated the emissions from the feeding site, although some sesquiterpenes were also emitted. Weevil feeding increased bark emission of monoterpenes by nearly 4-fold and sesquiterpenes by 7-fold. The influence of weevil damage on systemic monoterpene emissions from shoots was more profound. Several compounds were substantially induced, including linalool, β-phellandrene, limonene and 1,8-cineole. Sesquiterpenes contributed only 1.2% of the total foliage emission, but comprised eight different compounds including (E,E)-α-farnesene, β-bourbonene and (E)-β-farnesene. The total emission of monoterpenes and sesquiterpenes from shoots was respectively 2.8-fold and 2.9-fold higher in the pine weevil damaged plants than the undamaged plants. As many of the induced compounds are highly reactive in the atmosphere and form organic aerosol particles, our results suggest that conifers damaged by insects could become a more important source of secondary organic aerosols than healthy trees.  相似文献   

10.
Although considerable evidence has accumulated on the defensive activity of plant volatile organic compounds against pathogens and insect herbivores, less is known about the significance of volatile organic compounds emitted by plants under abiotic stress. Here, we report that green leaf volatiles (GLVs), which were previously shown to prime plant defences against insect herbivore attack, also protect plants against cold stress (4 °C). We show that the expression levels of several cold stress‐related genes are significantly up‐regulated in maize (Zea mays) seedlings treated with physiological concentrations of the GLV, (Z)‐3‐hexen‐1‐yl acetate (Z‐3‐HAC), and that seedlings primed with Z‐3‐HAC exhibit increased growth and reduced damage after cold stress relative to unprimed seedlings. Together, these data demonstrate the protective and priming effect of GLVs against cold stress and suggest an activity of GLVs beyond the activation of typical plant defence responses against herbivores and pathogens.  相似文献   

11.
Scots pine [ Pinus sylvestris L. (Pinaceae)] trees with four different seed origins were exposed to exogenous applications of the elicitor, methyl jasmonate (MeJA), for three consecutive years. We studied the effects of MeJA on needle chemistry (including monoterpenes, sesquiterpenes, and tricyclic resin acids), plant growth, and the performance of two diprionid sawflies, the European pine sawfly ( Neodiprion sertifer Geoffr.) and the common pine sawfly ( Diprion pini L.) (both Hymenoptera: Diprionidae). In general, foliar MeJA application affected the whole range of needle secondary chemistry with significantly higher concentrations of two monoterpenes, β-pinene and limonene, in particular. Furthermore, for some seed origins the growth rates of N. sertifer and D. pini larvae were lower on needles of MeJA-treated plants with either high total terpene or high resin acid concentrations. However, inconsistencies in diprionid sawfly performance within each studied Scots pine origin suggest genetic variance in needle secondary chemistry. The differences between selected seed origins and notably variable responses to MeJA application imply that adaptation of the seed to new conditions may have had an impact on secondary chemistry and, thus, on insect performance. Finally, our results suggest that modification of Scots pine defence by a low-concentration exogenous elicitor affects the production of terpenoids in the newly growing needles, leading to poorer pine sawfly performance in origins with high terpenoid content, while not harming the growth of Scots pine trees.  相似文献   

12.
Yan ZG  Wang CZ 《Phytochemistry》2006,67(1):34-42
Green leaf volatiles (GLVs), generally occurring C6 alcohols, aldehydes and acetates from plants, play an important role in plant-plant communication. These compounds induce intact plants to produce jasmonic acid, and induce defense-related gene expression and the release of volatile compounds. Here, we address wound-induced GLVs cause the release of acetylated derivatives and a terpenoid, (E)-4,8-dimethylnona-1,3,7-triene (DMNT) in intact maize, which may be a type of plant-plant interaction mediated by airborne GLVs. Upon exposure of intact maize seedlings to wound-induced GLVs, (Z)-3-hexenyl acetate was consistently the most abundant compound released. Exogenous application of individual alcohols and aldehydes mostly resulted in the release of corresponding acetate esters. C6-alcohols with a double bond between the second and third, or the third and fourth carbon atoms, C5- or C6-aldehydes, and (Z)-3-hexenyl acetate triggered the release of DMNT. When (Z)-3-hexenyl acetate and hexyl acetate were used to treat maize seedlings, they were recovered from the plants. These data demonstrated that: (1) apart from direct adsorption and re-release of acetate esters, absorption and conversion of exogenous alcohols and aldehydes into acetate esters occurred, and (2) DMNT was induced by a range of aldehydes and unsaturated alcohols.  相似文献   

13.
Volatiles emitted from the leaves of Lycopersicon esculentum at the two-, ten-leaf and anthesis periods were collected by a gas absorbing method and analyzed by gas chromatography (GC)-mass spectrometry. In total, 33 compounds of volaUles emitted from three developmental stage plants were separated and identified, and quantitatively analyzed by the internal standard addition method. All of the samples of volatile were found to be rich in monoterpenes and sesquiterpenes.β-phellandrene and caryophyllene predominated in the volatiles of the leaves of plants at the two- and ten-leaf stages. Furthermore, (E)-2-hexenal were the dominant components in the volatiles emitted from anthesis plants. The results of volatiles analyzed show that the compositions varied depending on the developmental stages. The volatiles emitted from crushed tomato leaves of plants at the anthesis stage had the most strongly inhibitory activity against the spore germination and hyphal growth of Botrytis cinerea and Fusarium oxysporum, followed by ten- and two-leaf plants. However, the activity of volatiles, emitted from the leaves of plants at the two-leaf stage, in inhibiting F. oxysporum was greater than B. cinerea.  相似文献   

14.
Certain species of Scrophularia (Scrophulariaceae), such as S. nodosa and S. umbrosa, are mainly pollinated by social wasps and are consequently described as wasp-flowers. Because plants attract their pollinators with the help of various floral cues, such as floral odour and/or optical cues, we have investigated the role of olfactory and visual floral signals responsible for wasp attraction in S. umbrosa. Using a combination of chemical (GC, GC-MS) and electrophysiological analyses (GC-EAD), we identified ten compounds in the complex floral odour bouquet that are detectable by the wasps' antennae. As in the wasp-flower Epipactis helleborine, we found so-called 'green leaf volatiles' (GLVs) in the floral odour; these GLVs are highly attractive to the wasps. GLVs, mostly six-carbon aldehydes, alcohols and acetates, and other volatile organic compounds (VOCs), are emitted by many plants infested with herbivores, e.g. caterpillars. In contrast to other investigated wasp-flowers, behavioural experiments have demonstrated that, in addition to the floral odour of S. umbrosa, visual cues are involved in pollinator attraction.  相似文献   

15.
The flowers of many plants emit volatile compounds as a means of attracting pollinators. We have previously shown that the strong, sweet fragrance of Clarkia breweri (Onagraceae), an annual plant native to California, consists of approximately 8 to 12 volatile compounds[mdash]three monoterpenes and nine benzoate derivatives (R.A. Raguso and E. Pichersky [1994] Plant Syst Evol [in press]). Here we report that the monoterpene alcohol linalool is synthesized and emitted mostly by petals but to a lesser extent also by the pistil and stamens. Two linalool oxides are produced and emitted almost exclusively by the pistil. These three monoterpenes are first discernible in mature unopened buds, and their tissue levels are highest during the first 2 to 3 d after anthesis. Levels of emission by the different floral parts throughout the life span of the flower were correlated with levels of these monoterpenes in the respective tissues, suggesting that these monoterpenes are emitted soon after their synthesis. Activity of linalool synthase, an enzyme that converts the ubiquitous C10 isoprenoid intermediate geranyl pyrophosphate to linalool, was highest in petals, the organ that emits most of the linalool. However, linalool synthase activity on a fresh weight basis was highest in stigma and style (i.e. the pistil). Most of the linalool produced in the pistil is apparently converted into linalool oxides. Lower levels (0.1%) of monoterpene emission and linalool synthase activity are found in the stigma of Clarkia concinna, a nonscented relative of C. breweri, suggesting that monoterpenes may have other functions in the flower in addition to attracting pollinators.  相似文献   

16.
Herbivore damage and mechanical injury to leaves can stimulate the emission of volatile compounds. It is well known that emission of these volatile organic compounds (VOC) from plants can influence interactions with pests and their natural enemies. In our experiment, we studied the VOC responses of Triticum aestivum cv. ‘Bombona’, Avena sativa cv. ‘Deresz’ and Hordeum vulgare cv. ‘Rastik’ under mechanical injury and/or adult cereal leaf beetle herbivory, Oelema melanopus (Coleoptera: Chrysomelidae). In the first part of our experiment, we confirmed that increased amounts of several green leaf volatiles (GLVs) and terpene VOC were released by tested cereal plants after leaf injury. The quantities of multiple induced VOC varied significantly between our tested cereals. When undamaged wheat, barley and oat plants were positioned near to mechanically injured or insect-damaged wheat plants, these neighbouring uninjured plants of all three test cereals also emitted significantly more VOC than control plants. The degree of VOC induction was significantly greater when an uninjured plant was closer to an injured wheat plant. This phenomenon may be useful for crop protection, as VOC manipulation may result in improved pest management and help reduce the use of harmful pesticides.  相似文献   

17.
The volatile profiles from flowers of Nicotiana sylvestris and N. suaveolens were investigated by means of dynamic headspace sampling and capillary gas chromatography. Under conditions of light/dark entrainment both species emitted phenylpropanoid-derived volatiles (e.g. benzyl alcohol, methyl benzoate) with maximum emission occurring during the dark period. Emission of these compounds was demonstrated to be circadian by continuance of rhythmicity under conditions of constant light and subsequent re-entrainment to a new light/dark cycle. In contrast, emission of the sesquiterpene hydrocarbon, caryophyllene, from N. sylvestris followed no apparent pattern. The emission of monoterpene hydrocarbons from flowers of N. suaveolens showed diurnal differences only under conditions of light/dark entrainment.  相似文献   

18.
The seasonal pattern of terpene content and emission by seven Mediterranean woody species was studied under field conditions. Emission rates were normalized at 30°C and 1000 μmol·m·s PFD (photosynthetic photon flux density). Bupleurum fruticosum, Pinus halepensis, and Cistus albidus stored large amounts of terpenes (0.01-1.77% [dry matter]) with maximum values in autumn and minimum values in spring. They emitted large amounts of terpenes (2-40 μg·g DM·h), but with no clear seasonal trend except for Cistus albidus, which had maximum values in spring and minimum values in autumn. The nonstoring species Arbutus unedo, Erica arborea, Quercus coccifera and Quercus ilex also emitted large amounts of terpenes (0-40 μg·g DM·h) and also tended to present maximum emission rates in spring, although this trend was significant only for A. unedo. At the seasonal scale, emission rates did not follow changes in photosynthetic rates; instead, they mostly followed changes in temperature. From autumn to spring, the least volatile monoterpenes such as limonene were emitted at highest rates, whereas the most volatile monoterpenes such as α-pinene and β-pinene were the most emitted in summer. The monoterpene emission rates represented a greater percentage of the photosynthetic carbon fixation in summer (from 0.51% in Arbutus unedo to 5.64% in Quercus coccifera) than in the rest of the seasons. All these seasonality trends must be considered when inventorying and modeling annual emission rates in Mediterranean ecosystems.  相似文献   

19.
Plant volatiles induced by herbivory are often used as olfactory cues by foraging herbivores and their natural enemies, and thus have potential for control of agricultural pests. Compared to chewing insects and mites, little is known about plant volatile production following herbivory by insects with piercing-sucking mouthparts. Here, we studied factors (insect life stage, gender, the role of salivary glands, and type of bioassay used for volatile induction) that influence the induction of plant volatiles by two agriculturally important hemipterans, Lygus hesperus and Nezara viridula. Feeding on intact cotton by virgin females of L. hesperus induced 2.6-fold greater volatile response compared to that induced by mated females, possibly due to increased feeding activity by virgin females. This plant volatile response was associated with elicitors present in the insect's salivary glands as well as to the degree of mechanical injury. Feeding injury by N. viridula females also increased volatile emissions in intact maize by approximately 2-fold compared to control plants. Maize seedlings injured by N. viridula emitted higher amounts of the monoterpene linalool, the sesquiterpenes (E)-beta-caryophyllene, alpha-trans-bergamotene, and (E,E)-beta-farnesene, and the homoterpene (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene, but not amounts of green leaf volatiles, compared to uninjured plants. Emissions from intact maize injured by adult males were lower than those emitted by adult females of the same age and did not differ from those emitted by uninjured plants. Similarly, feeding by virgin female N. viridula followed by excision led to 64% higher quantities of volatiles compared to untreated plants. Volatile emission in excised plants, however, was considerably greater than in intact plants, suggesting that careful consideration must be given to bioassay design in studies of herbivore-induced plant volatiles. Salivary gland extracts of N. viridula led to sesquiterpene emissions approximately 2.5-fold higher than for controls, although no significant differences were observed for green leaf volatiles, monoterpenes, and homoterpenes. These results indicate that L. hesperus and female N. viridula feeding induce volatile production in plants, and that volatile production is affected by gender and life stage of the bug. Although oviposition and mechanical injury by stylets may increase release of volatiles, elicitors from salivary glands of L. hesperus and N. viridula also seem to play a role in the emission of plant volatiles.  相似文献   

20.
Plants respond adaptively to herbivore stress in order to maintain fitness. Upon herbivore attack, plants emit blends of volatile organic compounds (VOCs) that differ from those that are constitutively emitted. These defense responses are typically specific to the identity of the attacking herbivore and often linked to the herbivore's feeding guild (e.g. chewing, phloem-feeding). Herbivores use plant volatiles to locate suitable host plants and changes in volatile emissions can affect host-plant location. Therefore, herbivores from separate feeding guilds can interact indirectly through the modulation of plant responses. In this study we tested how damage by an herbivore from one feeding guild affected the host-plant choice of an herbivore from a separate feeding guild, and vice versa. A chewing herbivore, the Colorado potato beetle (Leptinotarsa decemlineata), and a phloem feeding herbivore, the green peach aphid (Myzus persicae), were assayed in olfactometers to assess behavioral responses to odors emitted by potato plants (Solanum tuberosum) that were damaged by herbivores from the other feeding guild. Leptinotarsa decemlineata oriented more frequently towards undamaged plants compared to M. persicae damaged plants. Surprisingly, M. persicae preferred plants that were damaged by L. decemlineata, although previous studies had shown that they perform worse on these plants. Distinct differences were detected in the volatile profiles of herbivore-damaged and undamaged plants. Leptinotarsa decemlineata induced stronger volatile emissions compared to undamaged control plants, while M. persicae tended to suppress volatile emissions. These herbivores demonstrate contrasting induction of plant volatiles and behavioral responses. Exploring the nature of co-occurring herbivores and how they perceive potential hosts can play a significant role in understanding the ecological functions and community dynamics of plant plasticity and interactions with a variety of herbivores.  相似文献   

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