The Role of Fresh versus Old Leaf Damage in the Attraction of Parasitic Wasps to Herbivore-Induced Maize Volatiles

The odor produced by a plant under herbivore attack is often used by parasitic wasps to locate hosts. Any type of surface damage commonly causes plant leaves to release so-called green leaf volatiles, whereas blends of inducible compounds are more specific for herbivore attack and can vary considerably among plant genotypes. We compared the responses of naïve and experienced parasitoids of the species Cotesia marginiventris and Microplitis rufiventris to volatiles from maize leaves with fresh damage (mainly green leaf volatiles) vs. old damage (mainly terpenoids) in a six-arm olfactometer. These braconid wasps are both solitary endoparasitoids of lepidopteran larvae, but differ in geographical origin and host range. In choice experiments with odor blends from maize plants with fresh damage vs. blends from plants with old damage, inexperienced C. marginiventris showed a preference for the volatiles from freshly damaged leaves. No such preference was observed for inexperienced M. rufiventris. After an oviposition experience in hosts feeding on maize plants, C. marginiventris females were more attracted by a mixture of volatiles from fresh and old damage. Apparently, C. marginiventris has an innate preference for the odor of freshly damaged leaves, and this preference shifts in favor of a blend containing a mixture of green leaf volatiles plus terpenoids, after experiencing the latter blend in association with hosts. M. rufiventris responded poorly after experience and preferred fresh damage odors. Possibly, after associative learning, this species uses cues that are more directly related with the host presence, such as volatiles from host feces, which were not present in the odor sources offered in the olfactometer. The results demonstrate the complexity of the use of plant volatiles by parasitoids and show that different parasitoid species have evolved different strategies to exploit these signal

Mikroorganismen an Aesculus hippocastanum – olfaktorische Perspektive von Cameraria ohridella (Deschka & Dimic)

Since the 80s, the popular park- and city tree Aesculus hippocastanum has been infested with the leaf miner Cameraria ohridella (DESCHKA & DIMIC 1986). Additionally, the pathogenic fungi Guignardia aesculi causes leaf blotch disease and Erysiphe flexuosa causes powdery mildew on horse chestnuts. Often, all three diseases occur in parallel at the same leaves leading to a situation of competition. Moreover, recently some endophytic fungi were isolated from the leaf tissue of A. hippocastanum. In the present study, the volatile interaction between three trophic levels plant, insect, and fungi are discussed.Mikroorganismen an Aesculus hippocastanum – olfaktorische Perspektive von Cameraria ohridella (Deschka & Dimic) Seit den 80er Jahren wird die Gemeine Rosskastanie Aesculus hippocastanum durch den Minierer Cameraria ohridella (Lepidoptera, Gracillariidae) befallen. Außerdem wird die Rosskastanie durch den Blattbräunepilz Guignardia aesculi und den Mehltau Erysiphe flexuosa attackiert. Oft treten alle drei Organismen parallel am gleichen Blatt auf. Weiterhin konnten endophytische Pilze aus dem Blattgewebe isoliert werden. Im vorliegenden Beitrag wird die volatile Interaktion zwischen Pflanze, Mikroorganismen und Insekt diskutiert. Mit Hilfe der Gaschromatographie und gekoppelter Massenspektroskopie (GC-MS) wurden Duftproben gesunder und mit den pathogenen Pilzen G. aesculi und E. flexuosa gleichzeitig infizierter Blätter der Rosskastanie analysiert. Identifizierte Komponenten wurden elektrophysiologisch (EAG) an der Insektenantenne und in Verhaltensversuchen getestet. Mit den pathogenen Pilzen befallene Rosskastanienblätter geben 1-Octen-3-ol, 3-Octanon, ein Derivat von 2(5H)-Furanon, Nonanal und Decanal ab. C. ohridella war in der Lage, diese Substanzen zu detektieren. In Zweifachwahltests mit gesunden A. hippocastanum Zweigen reagierten Weibchen mit reduzierter Eiablage auf die Applikation von 1-Octen-3-ol, 3-Octanon, 2(5H)-Furanon und Decanal im Vergleich zur unbehandelten Kontrolle. Es ist bekannt, dass 1-Octen-3-ol und 3-Octanon von Pilzen selber emittiert werden. Nonanal und Decanal werden von Zellen, die nach Penetration durch Pilzhyphen unter oxidativem Stress stehen, produziert. Die Derivate von 2(5H)-Furanon wirken antimikrobiell und können auf einen Schutzmechanismus der Pflanze oder auf einen Konkurrenzmechanismus von Mikroorganismen um denselben Lebensraum hinweisen. Eine mögliche Erklärung wäre, dass diese Substanz von Endophyten zur Verteidigung des sie umgebenden Blattgewebes gegen die pathogenen Pilze produziert wird

Effects of virus infection on release of volatile organic compounds from insect-damaged bean, Phaseolus vulgaris

Insects can serve as important vectors of plant pathogens, especially viruses. Insect feeding on plants causes the systemic release of a wide range of plant volatile compounds that can serve as an indirect plant defense by attracting natural enemies of the herbivorous insect. Previous work suggests that the Mexican bean beetle (Epilachna varivestis) prefers to feed on plants infected by either of two viruses that it is known to transmit: Southern bean mosaic virus (SBMV) or Bean pod mottle virus (BPMV). A possible explanation for the preferred feeding on virus-infected tissues is that the beetles are attracted by volatile signals released from leaves. The purpose of this work was to determine whether volatile compounds from virus-infected plants are released differentially from those emitted by uninfected plants. To test the hypothesis, common bean plants (Phaseolus vulgaris cv. Black Valentine) were inoculated with either BPMV, SBMV, or a mixture of both viruses, and infected plants were compared to uninfected plants. An Ouchterlony assay was used with SBMVand BPMV-specific antisera to confirm the presence of virus in inoculated plants. RNA blot analysis was performed on tissue from each plant and indicated that a well-characterized defense gene, encoding phenylalanine ammonia-lyase (PAL), was not induced in systemic tissue following virus infection. Plant volatiles were collected—and analyzed via gas chromatography (GC)—from plants that were either undamaged or beetle-damaged. In undamaged plants, there were no measurable differences in profiles or quantities of compounds released by uninfected and virus-infected plants. After Mexican bean beetles were allowed to feed on plants for 48 h, injured plants released several compounds that were not released from undamaged plants. Lower quantities of volatile compounds were released from virus-infected plants suggesting that enhanced release of plant-derived volatile organic compounds is not the cause for attraction of Mexican bean beetles to virus-infected plants

Beeinflussen verschiedene Volatilenmuster die Wirtspflanzenwahl unterirdisch lebender Insekten?

Cockchafers of the genus Melolontha (Coleoptera: Scarabaeidae) can be severe pests in forestry, agriculture and horticulture. Gradation of the two most important species, the forest cockchafer M. hippocastani FABR. and the European cockchafer M. melolontha L., occurs currently in several parts of central Europe. Orientation behaviour of the adult beetles has been the focus of recent studies (REINECKE et al. 2002 a, b, 2005). However, especially the larvae are dreaded because their belowground damage is not visible directly after feeding. There are a lot of speculations about belowground living insects and their way of living, but until now there were not that many experimental investigations. A rather unknown topic is the orientation behaviour of soil living organisms, which is also subject of some publications (HORBER 1954, HAUSS & SCHÜTTE 1976, HASLER 1986, HIBBARD et al. 1994, JEWETT & BJOSTAD 1996, BERNKLAU & BJOSTAD 1998A, BERNKLAU & BJOSTAD 1998B, BERNKLAU et al. 2005).Zum besseren Verständnis der Orientierung und Fraßpräferenz von Maikäferlarven Melolontha hippocastani FABR. (Coleoptera: Scarabaeidae) im Boden wurden Wahltests durchgeführt. Den Larven von M. hippocastani wurden Karotten (Daucus carota ssp. sativus) und Eichenwurzeln (Quercus sp.) zur Auswahl angeboten. Die Duftstoffe von Karotte und Eichenwurzel wurden auf Aktivkohle gesammelt und mit Gaschromatographie – Massenspektroskopie untersucht (GC-MS). Unverletzte Karotten sowie Eichenwurzeln unterschieden sich in ihr n Volatilenmustern deutlich voneinander. Darüber hinaus konnten Unterschiede im Volatilenmuster unverletzter, mechanisch verletzter sowie angefressener Wurzeln nachgewiesen werden

Secondary Organic Aerosol Formation from Healthy and Aphid-Stressed Scots Pine Emissions.

One barrier to predicting biogenic secondary organic aerosol (SOA) formation in a changing climate can be attributed to the complex nature of plant volatile emissions. Plant volatile emissions are dynamic over space and time, and change in response to environmental stressors. This study investigated SOA production from emissions of healthy and aphid-stressed Scots pine saplings via dark ozonolysis and photooxidation chemistry. Laboratory experiments using a batch reaction chamber were used to investigate SOA production from different plant volatile mixtures. The volatile mixture from healthy plants included monoterpenes, aromatics, and a small amount of sesquiterpenes. The biggest change in the volatile mixture for aphid-stressed plants was a large increase (from 1.4 to 7.9 ppb) in sesquiterpenes-particularly acyclic sesquiterpenes, such as the farnesene isomers. Acyclic sesquiterpenes had different effects on SOA production depending on the chemical mechanism. Farnesenes suppressed SOA formation from ozonolysis with a 9.7-14.6% SOA mass yield from healthy plant emissions and a 6.9-10.4% SOA mass yield from aphid-stressed plant emissions. Ozonolysis of volatile mixtures containing more farnesenes promoted fragmentation reactions, which produced higher volatility oxidation products. In contrast, plant volatile mixtures containing more farnesenes did not appreciably change SOA production from photooxidation. SOA mass yields ranged from 10.8 to 23.2% from healthy plant emissions and 17.8-26.8% for aphid-stressed plant emissions. This study highlights the potential importance of acyclic terpene chemistry in a future climate regime with an increased presence of plant stress volatiles

An unbiased approach elucidates variation in (S)-(+)-linalool, a context-specific mediator of a tri-trophic interaction in wild tobacco

Plant volatile organic compounds (VOCs) mediate many interactions, and the function of common VOCs is especially likely to depend on ecological context. We used a genetic mapping population of wild tobacco, Nicotiana attenuata, originating from a cross of 2 natural accessions from Arizona and Utah, separated by the Grand Canyon, to dissect genetic variation controlling VOCs. Herbivory-induced leaf terpenoid emissions varied substantially, while green leaf volatile emissions were similar. In a field experiment, only emissions of linalool, a common VOC, correlated significantly with predation of the herbivore Manduca sexta by native predators. Using quantitative trait locus mapping and genome mining,we identified an (S)-(+)-linalool synthase (NaLIS). Genome resequencing, gene cloning, and activity assays revealed that the presence/absence of a 766-bp sequence in NaLIS underlies the variation of linalool emissions in 26 natural accessions. We manipulated linalool emissions and composition by ectopically expressing linalool synthases for both enantiomers, (S)-(+)- and (R)-(−)-linalool, reported to oppositely affect M. sexta oviposition, in the Arizona and Utah accessions.We used these lines to test ovipositingmoths in increasingly complex environments. The enantiomers had opposite effects on oviposition preference, but themagnitude of the effect depended strongly both on plant genetic background, and complexity of the bioassay environment. Our study reveals that the emission of linalool, a common VOC, differs by orders-of-magnitude among geographically interspersed conspecific plants due to allelic variation in a linalool synthase, and that the response of a specialist herbivore to linalool depends on enantiomer, plant genotype, and environmental complexity

Response of female Cydia molesta (Lepidoptera: Tortricidae) to plant derived volatiles

Peach shoot volatiles were attractive to mated female oriental fruit moth, Cydia molesta (Busck), in a dual choice arena. No preference was observed between leaf odours from the principle host plant, peach, and the secondary host plant, apple. Twenty-two compounds were identified in headspace volatiles of peach shoots using gas chromatography-mass spectrometry. Green leaf volatiles accounted for more than 50% of the total emitted volatiles. A bioassay-assisted fractionation using different sorbent polymers indicated an attractant effect of compounds with a chain length of 6-8 carbon atoms. The major compounds of this fraction were tested either singly or in combinations for behavioural response of females. Significant bioactivity was found for a three-component mixture of (Z)-3-hexen-1-yl acetate, (Z)-3-hexen-1-ol and benzaldehyde in a 4:1:1 ratio. This synthetic mixture elicited a similar attractant effect as the full natural blend from peach shoots as well as the bioactive fractio

Pollinator-attracting semiochemicals of the wasp-flower Epipactis helleborine

The orchid genus Epipactis is represented by 25 species in Europe (Richards 1982). Epipactis helleborine (L.) Crantz is the most common and widely distributed species of the genus (Wiefelspütz 1970), and is a prime example for wasp-flowers, because it is mainly pollinated by social wasps (Hymenoptera: Vespidae), like Vespula vulgaris and V. germanica (Müller 1873). Darwin (1888) already noticed that E. helleborine is almost exclusively ignored by bees and bumblebees, an observation that was confirmed in recent investigations (Keppert 2001). The flowers of E. helleborine show morphological, physiological and phenological adaptations to the visit and the pollination by Vespidae (Keppert 2001). They possess a reddish-brown or dirty purple coloration of the inflorescence (Keppert 2001), have relatively small, mostly bulbous blossoms with a broad entrance and bulbous widened, nectar-rich juice holders (Müller 1873, 1881; Schremmer 1962). Although there is much reported about wasp-pollinated flowers there is little known about the signals that are responsible for the attraction of wasps. Wiefelspütz (1970) proclaimed the statement that only the visual stimulus is responsible for the wasp attraction. Recently studies, however, assumed that odour is involved in the wasp attraction (Keppert 2001). Hölzler (2003) showed that the main attraction of the wasp-flower Epipactis for pollinators is its olfactory stimulus. It remains an unanswered question why E. helleborine flowers almost exclusively attracts social wasps, as opposed to bees and bumblebees. In this study we analysed the role of floral volatiles which are responsible for the specific attraction of social wasps. We supposed a mimicry-system in E. helleborine for the specific attraction of pollinators for the following reasons. So-called “green leaf volatiles” (GLVs) are emitted by plants while herbivorous insects, for example caterpillars, feed on them. GLVs thereby attract predators or parasitoids of the herbivorous insects (Dicke & Sabelis 1988; Turlings & al. 1990, 1995; Dicke & Vet 1999). Among the GLVs so far identified in former studies there are aldehydes, compounds that were also found in flower extracts of E. helleborine (Hölzler 2003). Therefore, we postulated that E. helleborine flowers produce GLVs in order to attract prey hunting social wasps for pollination. We performed bioassays and analysed flower odour gained to headspace-sampling using gas chromatography (GC), mass spectrometry (GC-MS) and gas chromatography coupled with electrophysiological analysis (GC-EAD) to investigate the hypothesis that E. helleborine flowers mimic “green leaf volatiles” (GLVs) to attract their pollinators.Die Orchideenart Epipactis helleborine gilt als typische Wespenblume. Die Blüten weisen Anpassungen an den Besuch und die Bestäubung durch soziale Faltenwespen (Hymenoptera: Vespidae) auf und werden häufig durch Vespula vulgaris und V. germanica bestäubt. In früheren Untersuchungen konnte gezeigt werden, dass olfaktorische Reize bei der Bestäuberanlockung eine übergeordnete Bedeutung vor optischen Reizen haben (Hölzler 2003). Die Frage, warum E. helleborine fast ausschließlich ihren optimalen Bestäuber, die soziale Faltenwespe, zur Bestäubung anlockt, und nicht auch auf andere Blütenbesucher attraktiv wirkt, ist noch unbeantwortet. Wir untersuchten die Hypothese, dass E. helleborine Blüten GLVs, die von Herbivoren befallenen Pflanzen abgegeben werden, nachahmen, um Beute jagende Wespen zur Bestäubung anzulocken. Dazu sammelten und analysierten wir Duftstoffe von Epipactis Blüten und mit Pieris-Raupen befallenen Kohl und identifizierten vier gemeinsam vorkommende GLVs. In Y-Rohrtests konnte die wespenanlockende Wirkung dieser Verbindungen nachgewiesen werden