Inhibition of feeding by a generalist insect due to increased volatile leaf terpenes under nitrate-limiting conditions

Nitrogen-limited plants ofHeterotheca subaxillaris accumulate greater quantities of leaf volatile terpenes than do nitrogen-rich plants. A series of feeding trials were performed to determine if such nitrate-limited plants are better defended against generalist-feeding insect herbivores. Soybean looper (Pseudoplusia includens) larvae were fed leaves fromH. subaxillaris rosettes grown under high and low nitrate supply regimes. Larval consumption, growth, and survival declined as the leaf volatile terpene content increased. Larval consumption and growth were enhanced by higher plant nitrate supply and with increasing leafage. The results suggest that the higher quantity of volatile terpenes in the leaves of nitrate-limited plants may better defend these leaves against generalist-feeding insects.     

Systemic Induction of Allelochemicals in Glanded and Glandless Isogenic Cotton by Spodoptera exigua Feeding

Induction of systemic resistance to feeding of beet armyworm, Spodoptera exigua, was investigated in two isogenic lines of Stoneville 213 cotton, Gossypium hirsutum, that differed in the presence of pigment glands. In laboratory bioassays, larvae strongly preferred to feed on glandless cotton plants when presented a choice between undamaged terminal leaves of undamaged glanded and glandless plants. Feeding damage inflicted by S. exigua larvae on the two oldest leaves of glanded plants seven days prior to feeding bioassays caused larvae to prefer by 33-fold the undamaged terminal foliage from undamaged plants compared to that from damaged plants. Feeding damage on glandless plants caused only a 2.6-fold greater preference for terminal foliage from undamaged plants over foliage from previously damaged plants. Extracts of terminal foliage from glanded cotton damaged seven days earlier had significantly greater quantities of terpenoid aldehydes (hemigossypolone, gossypol, and heliocides) than did foliage from undamaged glanded plants. Terpenoid aldehydes were undetectable in extracts of both undamaged and previously damaged glandless plants. The profile of volatile compounds collected from the headspace of mechanically damaged terminal leaves of undamaged glanded and glandless plants differed. Both cotton isolines released large quantities of lipoxygenase products (hexenyl alcohols, acetates, and butyrates), but glandless plants released only small amounts of mono- and sesquiterpenes compared to glanded plants. Glandless plants damaged seven days prior to volatile collection released significantly greater quantities of lipoxygenase products, -ocimene, and - and -farnesene than did undamaged glandless plants. Previously damaged glanded plants released significantly greater quantities of all mono- and sesquiterpenes and hexenyl acetates and butyrates, but not alcohols. The relative importance of volatile compounds versus terpenoid aldehydes in induced feeding deterrence in cotton to S. exigua larvae is still unclear.     

Cyanide in the Chemical Arsenal of Garlic Mustard, <Emphasis Type="Italic">Alliaria petiolata</Emphasis>

Cyanide production has been reported from over 2500 plant species, including some members of the Brassicaceae. We report that the important invasive plant, Alliaria petiolata, produces levels of cyanide in its tissues that can reach 100 ppm fresh weight (FW), a level considered toxic to many vertebrates. In a comparative study, levels of cyanide in leaves of young first-year plants were 25 times higher than in leaves of young Arabidopsis thaliana plants and over 150 times higher than in leaves of young Brassica kaber, B. rapa, and B. napus. In first-year plants, cyanide levels were highest in young leaves of seedlings and declined with leaf age on individual plants. Leaves of young plants infested with green peach aphids (Myzus persicae) produced just over half as much cyanide as leaves of healthy plants, suggesting that aphid feeding led to loss of cyanide from intact tissues before analysis, or that aphid feeding inhibited cyanide precursor production. In a developmental study, levels of cyanide in the youngest and oldest leaf of young garlic mustard plants were four times lower than in the youngest and oldest leaf of young Sorghum sudanense (cv. Cadan 97) plants, but cyanide levels did not decline in these leaves with plant age as in S. sudanense. Different populations of garlic mustard varied moderately in the constitutive and inducible expression of cyanide in leaves, but no populations studied were acyanogenic. Although cyanide production could result from breakdown products of glucosinolates, no cyanide was detected in vitro from decomposition of sinigrin, the major glucosinolate of garlic mustard. These studies indicate that cyanide produced from an as yet unidentified cyanogenic compound is a part of the battery of chemical defenses expressed by garlic mustard.     

Environmental and Developmental Regulation of Trypsin Inhibitor Activity in Brassica napus

We examined several environmental and developmental influences on trypsin inhibitor (TI) activity in leaves of young Brassica napus seedlings in a series of greenhouse experiments. In seedlings of B. napus cv. Westar, TI activity is constitutively present and exhibits a rise then fall through time in the first true leaves of young plants. TI activity is induced by wounding in the first true leaves, but the degree of induction is relatively insensitive to the degree of wounding over a gradient of 5–15% of leaf area damage. TI activity is enhanced in first true leaves of plants in which the cotyledons have been wounded relative to plants in which the cotyledons have not been wounded. TI activity is also enhanced in the second true leaves on plants in which the first true leaves have been wounded. The degree of systemic induction in second true leaves declines additively with plant age, but local induction in the first true leaves is not affected by age. In B. napus cv. Gido, TI activity is constitutively present but is not locally wound-inducible in first true leaves of young plants exposed to the same wounding gradient as cv. Westar. In unwounded plants at the six-leaf stage, TI activity is higher in second true leaves than in fifth true leaves, indicating that TI activity is developmentally regulated in this cultivar.     

Attraction of adult sweet potato weevils,Cylas formicarius elegantulus (Summers), (Coleoptera: Curculionidae), to sweet potato leaf and root volatiles

A dual-choice olfactometer was developed to study the responses of sweet potato weevils,Cylas formicarius elegantulus (Summers), to volatiles from the sweet potato,Ipomoea batatas (L.) Lam. Both males and females were attracted by volatiles from sweet potato leaves and a methylene chloride leaf extract. Females, but not males, responded to volatiles from storage roots and a methylene chloride root extract. Leaves and storage roots from four sweet potato cultivars (Centennial, Jewel, Resisto, and Regal) were attractive to female weevils; however, the attractant response varied with cultivar. GC profiles from leaf and root extracts, and GC-MS analysis of leaf extract, for Jewel cultivar enabled the volatile peaks to be identified as sesquiterpenes.     

Citrus Leaf Volatiles as Affected by Developmental Stage and Genetic Type

Major volatiles from young and mature leaves of different citrus types were analyzed by headspace-solid phase microextraction (HS-SPME)-GC-MS. A total of 123 components were identified form nine citrus cultivars, including nine aldehydes, 19 monoterpene hydrocarbons, 27 oxygenated monoterpenes, 43 sesquiterpene hydrocarbons, eight oxygenated sesquiterpenes, two ketones, six esters and nine miscellaneous. Young leaves produced higher amounts of volatiles than mature leaves in most cultivars. The percentage of aldehyde and monoterpene hydrocarbons increased, whilst oxygenated monoterpenes and sesquiterpenes compounds decreased during leaf development. Linalool was the most abundant compound in young leaves, whereas limonene was the chief component in mature ones. Notably, linalool content decreased, while limonene increased, during leaf development in most cultivars. Leaf volatiles were also affected by genetic types. A most abundant volatile in one or several genotypes can be absent in another one(s), such as limonene in young leaves of lemon vs. Satsuma mandarin and β-terpinene in mature leaves of three genotypes vs. the other four. Compositional data was subjected to multivariate statistical analysis, and variations in leaf volatiles were identified and clustered into six groups. This research determining the relationship between production of major volatiles from different citrus varieties and leaf stages could be of use for industrial and culinary purposes.     

Distribution and Differential Expression of (E)-4,8-Dimethyl-1,3,7-Nonatriene in Leaf and Floral Volatiles of Magnolia and Liriodendron Taxa

Analyses of volatiles emitted from artificially damaged leaves attached to branches of seven Magnolia taxa revealed the presence of (Z)-3-hexenyl acetate, (Z)-3-hexenol (the green odor compounds), and several mono- and sesquiterpenes, e.g., (Z)- and (E)-β-ocimene and caryophyllene. An herbivore-induced leaf volatile, (E)-4,8-dimethyl-1,3,7-nonatriene, known as a predator attractant in agricultural plants, was emitted 4–6 hr after leaves were damaged in M. hypoleuca. The damaged leaves of M. grandiflora, however, immediately released (E)-4,8-dimethyl-1,3,7-nonatriene. Undamaged leaves of Magnolia species examined did not emit volatile compounds. In addition, detached flowers of six Magnolia taxa and Liriodendron tulipifera also emit (E)-4,8-dimemyl-1,3,7-nonatriene as a floral volatile (up to 30% in some species); the chemical was also emitted from the intact flowers of M. heptapeta and M. salicifolia.     

Detrimental effects ofCinchona leaf alkaloids on larvae of the polyphagous insectSpodoptera exigua

YoungCinchona ledgeriana plants contain two types of alkaloid: indole alkaloids in the leaves and quinoline alkaloids in the root. FromCinchona leaves, a crude alkaloid extract was made, containing the cinchophylline type of indole alkaloids and a small amount of 5-methoxytryptamine. The leaf alkaloid extract exerted a strong detrimental effect on the growth of larvae of the polyphagous beet armyworm,Spodoptera exigua (Lepidoptera). Feeding of larvae on an artificial diet containing the leaf alkaloids at the same concentrations as those found in the plant resulted in significant growth reduction, retardation in development, and mortality of the larvae. Cinchophyllines are composed of 5-methoxytryptamine coupled to a corynantheal unit. When incorporated into the artificial diet, 5-methoxytryptamine alone had no effect on the 5.exigua larvae. Corynantheal, however, had a strong detrimental effect on growth of the larvae, its effect being comparable to that of the leaf alkaloid extract. In contrast to the indole-type leaf alkaloids, the quinolinetypeCinchona root alkaloids did not affect growth and development of the larvae. These results suggest that the indole-type alkaloids, which inCinchona plants are present at the highest concentrations in the young, vulnerable leaflets, are involved in the chemical defense of the plant against herbivorous insects.     

Foliar Terpene Chemotypes and Herbivory Determine Variation in Plant Volatile Emissions

Plants that synthesize and store terpenes in specialized cells accumulate large concentrations of these compounds while avoiding autotoxicity. Stored terpenes may influence the quantity and profile of volatile compounds that are emitted into the environment and the subsequent role of those volatiles in mediating the activity of herbivores. The Australian medicinal tea tree, Melaleuca alternifolia, occurs as several distinct terpene chemotypes. We studied the profile of its terpene emissions to understand how variations in stored foliar terpenes influenced emissions, both constitutive and when damaged either by herbivores or mechanically. We found that foliar chemistry influenced differences in the composition of terpene emissions, but those emissions were minimal in intact plants. When plants were damaged by herbivores or mechanically, the emissions were greatly increased and the composition corresponded to the constitutive terpenes and the volatility of each compound, suggesting the main origin of emissions is the stored terpenes and not de novo biosynthesized volatiles. However, herbivores modified the composition of the volatile emissions in only one chemotype, probably due to the oxidative metabolism of 1,8-cineole by the beetles. We also tested whether the foliar terpene blend acted as an attractant for the specialized leaf beetles Paropsisterna tigrina and Faex sp. and a parasitoid fly, Anagonia zentae. None of these species responded to extracts of young leaves in an olfactometer, so we found no evidence that these species use plant odor cues for host location in laboratory conditions.     

Chemical Defense in Larval Tortoise Beetles: Essential Oil Composition of Fecal Shields of Eurypedus nigrosignata and Foliage of its Host Plant, Cordia curassavica

Larvae of the tortoise beetle Eurypedus nigrosignata construct fecal shields using cast skins and fecal strands. Survival of larvae with intact shields was higher in the field than for larvae with shields removed. In the laboratory, E. nigrosignata feculae had a deterrent effect on feeding in the ant Myrmica rubra as did an extract of the host plant, Cordia curassavica. Three chemical types were identified in the host-plant foliage and were named -terpinene, -pinene, and sabinene, depending on their mono- and sesquiterpene composition. This is the first report of lower terpenes (essential oils) in foliage of Cordia. Fecal shields of E. nigrosignata displayed the same terpene pattern as larval host-plant leaves. The absolute concentration of mono- and sesquiterpenes in the dorsal fecal shield depended on the plant chemical type and tended to decrease with larval age. No oxidation or detoxification products of ingested terpenes were detected in the larval fecula, indicating that the chemical composition of the larval fecal shield is influenced primarily by the host-plant secondary chemistry.     

Volatiles Emission Patterns in Poplar Clones Varying in Response to Ozone

The volatiles emitted from young and old leaves of two poplar clones (Populus deltoides x maximowiczii, Eridano, and P. x euramericana, I-214) were sampled after exposure to ozone (80 ppb, 5 h?d(-1), for 10 consecutive days) by solid phase microextraction and characterized by GC-MS. Only mature leaves of the ozone-sensitive Eridano clone developed necrosis in response to ozone exposure, and their membrane integrity was significantly affected by ozone (+86 and +18 % of levels of thiobarbituric acid reactive substances in mature and young leaves). The headspace of the poplar clones studied here contained mono- and sesquiterpenes, both hydrocarbons and oxygenated ones in Eridano, and only hydrocarbons in the clone I-214. Furthermore, some non-terpenes, such as C(9)-C(15) straight-chain aldehydes and C(12)-C(16) saturated and unsaturated aliphatic hydrocarbons, were detected. Other common non-terpene volatiles were oxygenated aliphatic compounds, mainly C(6)-alcohols and their acetates. Ozone exposure induced a strong change in volatile profiles, depending on clones and leaf age. Regardless of leaf age, in clone I-214, quantities of oxygenated monoterpenes tended to increase after ozone exposure, however, "O(3) x leaf age" was not significant. In clone Eridano, increases were observed in emissions of hydrocarbons and oxygenated sesquiterpenes in response to ozone treatment. (Z)-3-Hexen-1-ol and (Z)-3-hexenol acetate were present in traces in the headspace of untreated Eridano mature leaves, but quantities slightly increased after ozone treatment. Quantities of non-terpene oxygenated compounds dropped in the headspace of young leaves of both clones (-24 and -44 % in Eridano and I-214) and also in mature ones of I-214 (-50 %) after ozone exposure. Similarly, quantities of non-terpene hydrocarbons in the emissions from mature leaves of both clones (-58 and -49 %, respectively) decreased, while these compounds increased in young leaves of Eridano (+83 %). We suggest that the resistance of the poplar clone I-214 to O(3) is achieved by: i) monoterpenes constitutively present in young leaves and ii) increase of monoterpene content induced by O(3) in mature leaves.     

Cotton Plant, Gossypium hirsutum L., Defense in Response to Nitrogen Fertilization

Plants respond to insect herbivory by producing dynamic changes in an array of defense-related volatile and nonvolatile secondary metabolites. A scaled response relative to herbivory levels and nutrient availability would be adaptive, particularly under nutrient-limited conditions, in minimizing the costs of expressed defensive pathways and synthesis. In this study, we investigated effects of varying nitrogen (N) fertilization (42, 112, 196, and 280 ppm N) on levels of cotton plant (Gossypium hirsutum) phytohormones [jasmonic acid (JA) and salicylic acid (SA)], terpenoid aldehydes (hemigossypolone, heliocides H₁, H₂, H₃, and H₄), and volatile production in response to beet armyworm (Spodoptera exigua) herbivory. Additional bioassays assessed parasitoid (Cotesia marginiventris) host-searching success in response to cotton plants grown under various N fertilizer regimes. At low N input (42 ppm N), herbivore damage resulted in significant increases in local leaf tissue concentrations of JA and volatiles and in systemic accumulation of terpenoid aldehydes. However, increased N fertilization of cotton plants suppressed S. exigua-induced plant hormones and led to reduced production of various terpenoid aldehydes in damaged mature leaves and undamaged young leaves. While increased N fertilization significantly diminished herbivore-induced leaf volatile concentrations, the parasitism of S. exigua larvae by the parasitoid C. marginiventris in field cages did not differ among N treatments. This suggests that, despite significant N fertilization effects on herbivore-induced plant defenses, at short range, the parasitoids were unable to differentiate between S. exigua larvae feeding on physiologically different cotton plants that share large constitutive volatile pools releasable when damaged by herbivores.     

Systemic Induction of Terpenoid Aldehydes in Cotton Pigment Glands by Feeding of Larval Spodoptera exigua

Pigment glands in cotton contain terpenoid aldehydes that are toxic and deterrent to feeding of several generalist lepidopteran insects. We hypothesized that previously observed systemically induced feeding deterrence may be associated with pigment glands. We conducted experiments to determine the dynamics and chemical nature of inducible feeding deterrents in leaves of cotton, Gossypium hirsutum L, to larvae of the beet armyworm, Spodoptera exigua. Production and/or filling of pigment glands was influenced by physiological age of Deltapine 90 cotton plants. In undamaged plants, successively formed leaves contained more pigment glands, up to the seventh or eighth true-leaf developmental stage. Feeding choice tests conducted one or seven days after initial feeding damage revealed that third instars of S. exigua consumed more of the two youngest leaves from control cotton plants than from plants whose two oldest leaves had been fed on previously for 24 hr by S. exigua. The preference for leaves from control plants was significant one day after initial damage and highly significant seven days after damage. Consumption of mature foliage (leaf immediately above initially damaged leaves) from control plants and damaged plants did not differ. More pigment glands were counted on the youngest leaf of damaged plants than on the youngest leaf of control plants one day after initial damage. HPLC analysis revealed greater amounts of hemigossypolone, heliocides 1 and 2 (H₁ and H₂), and total terpenoid aldehydes per gland in young foliage of damaged plants than control plants one day after initial injury. By seven days after initial injury, greater quantities of hemigossypolone and all heliocides except H₄ were detected in young foliage from damaged plants compared to control plants. Concentrations of H₁ per gland in young leaves from damaged plants increased the most of all terpenoid aldehydes measured (3.4× the amount found in leaves from control plants). Mature leaves from damaged plants did not contain more terpenoid aldehydes than mature leaves from control plants. We suggest that systemically induced feeding deterrence to S. exigua in young leaves of glanded cotton was due to increased amounts of terpenoid aldehydes in pigment glands.     

Aphid and Plant Volatiles Induce Oviposition in an Aphidophagous Hoverfly

Episyrphus balteatus DeGeer (Diptera, Syrphidae) is an abundant and efficient aphid-specific predator. We tested the electroantennographic (EAG) response of this syrphid fly to the common aphid alarm pheromone, (E)-β-farnesene (EβF), and to several plant volatiles, including terpenoids (mono- and sesquiterpenes) and green leaf volatiles (C6 and C9 alcohols and aldehydes). Monoterpenes evoked significant EAG responses, whereas sesquiterpenes were inactive, except for the aphid alarm pheromone (EβF). The most pronounced antennal responses were elicited by six and nine carbon green leaf alcohols and aldehydes [i.e., (Z)-3-hexenol, (E)-2-hexenol, (E)-2-hexenal, and hexanal]. To investigate the behavioral activity of some of these EAG-active compounds, E. balteatus females were exposed to R-(+)-limonene (monoterpene), (Z)-3-hexenol (green leaf alcohol), and EβF (sesquiterpene, common aphid alarm pheromone). A single E. balteatus gravid female was exposed for 10 min to an aphid-free Vicia faba plant that was co-located with a semiochemical dispenser. Without additional semiochemical, hoverfly females were not attracted to this plant, and no oviposition was observed. The monoterpene R-(+)-limonene did not affect the females’ foraging behavior, whereas (Z)-3-hexenol and EβF increased the time of flight and acceptance of the host plant. Moreover, these two chemicals induced oviposition on aphid-free plants, suggesting that selection of the oviposition site by predatory hoverflies relies on the perception of a volatile blend composed of prey pheromone and typical plant green leaf volatiles.     

Phenolic acids affect photosynthesis and protein synthesis by isolated leaf cells of velvet-leaf

The effects ofp-coumaric, ferulic, chlorogenic, and vanillic acids on photosynthesis and protein synthesis by isolated leaf cells of velvetleaf (Abutilon theophrasti Medik) were investigated. Photosynthesis and protein synthesis were measured in cell suspensions by the incorporation of¹⁴CO₂ and [¹⁴C]leucine, respectively. None of the tested phenolic acids except vanillic reduced photosynthesis by more than 50% at the highest concentration and 30 min of incubation. At 100M concentrations and 60-min incubation periods,p-coumaric, ferulic, chlorogenic, and vanillic acids inhibited photosynthesis by 33, 37, 57, and 65%, respectively. Ferulic acid was the most inhibitory to protein synthesis and reduced the incorporation of [¹⁴C]leucine by 50% at about 1.0M after 60 min of incubation. At the highest concentrations tested in this study, vanillic and ferulic acids were inhibitory to photosynthesis and protein synthesis, respectively, whereas chlorogenic andp-coumaric acids did not inhibit either physiological process. The maximum inhibition of protein synthesis by chlorogenic acid was 19% and by vanillic acid was 28% at 100M concentrations. Chlorogenic, vanillic, andp-cou-maric acids at 0.1M caused increased protein synthesis over the untreated control. Overall, photosynthesis was more sensitive than protein synthesis to the four phenolic acids tested.Florida Agricultural Experiment Station Journal Series No. 9228.     

Systemically Induced Plant Volatiles Emitted at the Time of “Danger”

Feeding by Pieris brassicae caterpillars on the lower leaves of Brussels sprouts (Brassica oleracea var. gemmifera) plants triggers the release of volatiles from upper leaves. The volatiles are attractive for a natural antagonist of the herbivore, the parasitoid Cotesia glomerata. Parasitoids are attracted only if additional damage is inflicted on the systemically induced upper leaves and only after at least three days of herbivore feeding on the lower leaves. Upon termination of caterpillar feeding, the systemic signal is emitted for a maximum of one more day. Systemic induction did not occur at low levels of herbivore infestation. Systemically induced leaves emitted green leaf volatiles, cyclic monoterpenoids, and sesquiterpenes. GC-MS profiles of systemically induced and herbivore-infested leaves did not differ for most compounds, although herbivore infested plants did emit higher amounts of green leaf volatiles. Emission of systemically induced volatiles in Brussels sprouts might function as an induced defense that is activated only when needed, i.e., at the time of caterpillar attack. This way, plants may adopt a flexible management of inducible defensive resources to minimize costs of defense and to maximize fitness in response to unpredictable herbivore attack.     

Volatile Emissions from an Odorous Plant in Response to Herbivory and Methyl Jasmonate Exposure

Induced volatile terpenes have been commonly reported among diverse agricultural plant species, but less commonly investigated in odorous plant species. Odorous plants synthesize and constitutively store relatively large amounts of volatiles, and these may play a role in defense against herbivores. We examined the effect of herbivory and methyl jasmonate (MeJA) exposure on the release of volatile organic compounds (VOCs) in the marsh elder, Iva frutescens, which contains numerous constitutive VOCs, mainly mono- and sesquiterpenes. Our specific goal was to test for the presence of inducible VOCs in a naturally occurring plant already armed with VOCs. The abundant, native specialist leaf beetle Paria aterrima was used in herbivore induction trials. VOCs were sampled from herbivore wounded and unwounded, and from MeJA treated and untreated I. frutescens. Total VOC emissions were significantly greater in response to herbivory and MeJA treatment compared to unwounded controls. Herbivore wounding caused a substantial shift in the emission profile (42 VOCs from wounded, compared to 8 VOCs from unwounded I. frutescens), and MeJA had a similar yet less substantial influence on the emission pattern (28 VOCs from MeJA treated compared to 8 VOCs from untreated I. frutescens). Constitutive VOC emissions predominated, but some VOCs were detected only in response to herbivory and MeJA treatment, suggesting de novo synthesis. Several VOCs exhibited a delayed emission profile in contrast to the rapid release of constitutive VOCs, and principal components analysis revealed they were not associated with constitutive emissions. While I. frutescens contains many constitutive VOCs that are released immediately in response to herbivory, it also produces novel VOCs in response to feeding by the specialist P. aterrima and MeJA treatment.     

The Mono - and Sesquiterpene Content of Aphid-Induced Galls on Pistacia palaestina is Not a Simple Reflection of Their Composition in Intact Leaves

Pistacia palaestina Boiss. (Anacardiaceae), a sibling species of P. terebinthus also known as turpentine tree or terebinth tree, is common in the Levant region. The aphid Baizongia pistaciae L. manipulates the leaves of the plant to form large galls, which provide both food and protection for its developing offspring. We analyzed the levels and composition of mono-and sesquiterpenes in both leaves and galls of ten naturally growing trees. Our results show that monoterpene hydrocarbons are the main constituents of P. palaestina leaves and galls, but terpene levels and composition vary among trees. Despite this inter-tree variation, terpene levels and compositions in galls from different trees resemble each other more than the patterns displayed by leaves from the same trees. Generally, galls contain 10 to 60 fold higher total terpene amounts than leaves, especially of the monoterpenes α-pinene and limonene. Conversely, the leaves generally accumulate more sesquiterpenes, in particular E-caryophyllene, germacrene D and δ-cadinene, in comparison to galls. Our results clearly show that the terpene pattern in the galls is not a simple reflection of that of the leaves and suggest that aphids have a strong impact on the metabolism of their host plant, possibly for their own defense.     

Influence of Feeding and Oviposition by Phytophagous Pentatomids on Photosynthesis of Herbaceous Plants

Feeding by herbivorous insects may change photosynthetic activity of host plants. We studied how feeding and oviposition by herbivorous stink bugs, Murgantia histrionica and Nezara viridula (Heteroptera: Pentatomidae), affect photosynthetic parameters of Brassica oleracea (savoy cabbage) and Phaseolus vulgaris (French bean). First, we measured photosynthetic gas exchange, chlorophyll fluorescence imaging, and emission of induced volatile organic compounds (VOC) immediately after feeding and during a post-feeding period. Photosynthesis decreased rapidly and substantially in B. oleracea and P. vulgaris infested by feeding bugs. Stomatal conductance did not decrease proportionally with photosynthesis; instead, inhibition of photosynthesis likely was due to a reduced diffusion of CO₂ in the mesophyll. We also measured the impact of oviposition per se and oviposition associated with feeding on photosynthetic parameters. A surprisingly large inhibition of photosynthesis was detected in cabbage leaves in response to oviposition by M. histrionica, even when oviposition was not associated with feeding activity. High resolution chlorophyll fluorescence imaging revealed that the damage to photochemistry caused by feeding and oviposition was restricted to the attacked areas. By contrast, the photochemical yield increased temporarily in the unaffected areas of the attacked leaves, indicating the onset of a compensatory response. Measurement of volatile organic compounds (VOC) revealed that feeding-damaged plants did not emit detectable amounts of VOC, indicating cellular damage (methanol and green leaf volatiles). However, feeding by M. histrionica induced emission of mono- and sesquiterpenes in savoy cabbage leaves. The different time-course of the induction of these two classes of terpenes may reflect the induction of two different biosynthetic pathways and indicate different roles of these terpenoids in tritrophic interactions.     

Electroantennogram responses ofCampoletis sonorensis (hymenoptera: Ichneumonidae) to chemicals in cotton (Gossypium hirsutum L.)

Combined gas chromatography-electroantennogram (GC-EAG) recording ofCampoletis sonorensis (Cameron) responses to cotton plant volatile chemicals was performed.C. sonorensis antennal olfactory receptors respond differentially to green leaf, mono-, and sesquiterpene chemicals that have been identified previously in cotton. EAG depolarizations to green leaf chemicals were greater than to terpenes.