Fertility,Root Reserves and the Cost of Inducible Defenses in the Perennial Plant <Emphasis Type="Italic">Solanum carolinense</Emphasis>

We examined the relationship between internal resources (root reserves), external resources (soil fertility), and allocation to defense vs. growth in the clonal, perennial herb Solanum carolinense. In a short-term (9 d) greenhouse experiment, plants were treated once with jasmonic acid (JA) to determine if polyphenols and glycoalkaloids were inducible by simulated herbivory. In a longer-term (4 wk) greenhouse experiment, we measured the cost, in terms of growth, of treatment with JA every 3 d, to determine if the induced response was due more to carbon limitation or nitrogen limitation. We manipulated the resources available to the plants by varying soil fertility and the size of root cuttings from which plants were grown, and assessed how different resource levels affected the growth and production of polyphenols and alkaloids under JA treatment or control conditions. In the short term, JA increased the concentration of polyphenols in both above- and belowground plant parts, as well as alkaloid concentrations in the roots. In the long term, the only significant secondary chemistry response to JA was an increased polyphenol concentration in above ground tissues. The total amount of polyphenols produced was the same for JA and control plants, indicating that the higher concentration was a result of the lower biomass of treated plants. In contrast, alkaloid concentrations in plants treated with JA for 4 wk did not differ from controls, but JA-treated plants contained lower total amounts of alkaloids in above ground tissues, as a result of decreased growth. Fertilizer level and root cutting size had effects on growth and the production of secondary compounds and influenced the cost of induction. Plants grown under high fertility had a greater reduction in growth in response to JA than plants grown under low fertility, indicating a greater trade-off between growth and defense for high fertility plants. Plants from larger root cuttings grew bigger without any reduction in the concentration of polyphenols and alkaloids. We demonstrated that the phenotype of S. carolinense was plastic in response to simulated herbivory, fertility level, and root cutting size, and that there was a significant growth cost to induction that varied with the environment and appears to be due in large part to the allocation of limited carbon reserves.     

Exogenous Application of Jasmonic Acid Induces Volatile Emissions in Rice and Enhances Parasitism of <Emphasis Type="Italic">Nilaparvata lugens</Emphasis> Eggs by theParasitoid <Emphasis Type="Italic">Anagrus nilaparvatae</Emphasis>

Jasmonate signaling pathway plays an important role in induced plant defense against herbivores and pathogens, including the emission of volatiles that serve as attractants for natural enemies of herbivores. We studied the volatiles emitted from rice plants that were wounded and treated with jasmonic acid (JA) and their effects on the host-searching behavior of the rice brown planthopper, Nilaparvata lugens (Stål), and its mymarid egg parasitoid Anagrus nilaparvatae Pang et Wang. Female adults of N. lugens significantly preferred to settle on JA-treated rice plants immediately after release. The parasitoid A. nilaparvatae showed a similar preference and was more attracted to the volatiles emitted from JA-treated rice plants than to volatiles from control plants. This was also evident from greenhouse and field experiments in which parasitism of N. lugens eggs by A. nilaparvatae on plants that were surrounded by JA-treated plants was more than twofold higher than on control plants. Analyses of volatiles collected from rice plants showed that JA treatment dramatically increased the release of volatiles, which included aliphatic aldehydes and alcohols, monoterpenes, sesquiterpenes, methyl salicylate, n-heptadecane, and several as yet unidentified compounds. These results confirm an involvement of the JA pathway in induced defense in rice plants and demonstrate that the egg parasitoid A. nilaparvatae exploits plant-provided cues to locate hosts. We explain the use of induced plant volatiles by the egg parasitoid by a reliable association between planthopper feeding damage and egg presence.     

Divergence in Structure and Activity of Phenolic Defenses in Young Leaves of Two Co-Occurring <Emphasis Type="Italic">Inga</Emphasis> Species

The leaves of tropical forest trees are most likely to suffer herbivore damage during the period of expansion. Herbivore selection on young leaves has given rise to a variety of leaf developmental strategies and age-specific chemical defense modes. We are studying correlations between leaf developmental types and chemical defenses in the Neotropical genus Inga. We have characterized defense metabolites in Inga goldmanii and Inga umbellifera, two species that co-occur in the lowland moist forest of Panama. These congeners have markedly different young-leaf developmental phenotypes but suffer approximately equal rates of herbivory. Bioassays of whole and fractionated leaf extracts using larvae of Heliothis virescens show that I. goldmanii chemical defenses are nearly three times more inhibitory than those of I. umbellifera. In both species, most of the inhibitory activity resides in complex mixtures of monomeric and polymeric flavan-3-ols. This group comprises >30% of young leaf dry weight in both I. goldmanii and I. umbellifera. The species’ phenolic chemistry differs markedly, however, both in the structure of the monomeric units and in the distribution of polymer sizes. The differences in chemical structure have pronounced effects on their bioactivities, with I. goldmanii flavans being twice as inhibitory to H. virescens larvae as I. umbellifera flavans, and more than three times more efficient at protein binding. Given the extraordinarily high polyphenol concentrations that are found in the young leaves of these species, protein precipitation could be an important mechanism of growth inhibition. Nevertheless, our data show that another mode of phenolic action, possibly oxidative stress, occurs simultaneously.     

Effects of Root Isoquinoline Alkaloids from <Emphasis Type="Italic">Hydrastis canadensis</Emphasis> on <Emphasis Type="Italic">Fusarium oxysporum</Emphasis> Isolated from <Emphasis Type="Italic">Hydrastis</Emphasis> Root Tissue

Goldenseal (Hydrastis canadensis L.) is a popular medicinal plant distributed widely in North America. The rhizome, rootlets, and root hairs produce medicinally active alkaloids. Berberine, one of the Hydrastis alkaloids, has shown antifungal activity. The influence of a combination of the major Hydrastis alkaloids on the plant rhizosphere fungal ecology has not been investigated. A bioassay was developed to study the effect of goldenseal isoquinoline alkaloids on three Fusarium isolates, including the two species isolated from Hydrastis rhizosphere. The findings suggest that the Hydrastis root extract influences macroconidia germination, but that only the combined alkaloids—berberine, canadine, and hydrastine—appear to synergistically stimulate production of the mycotoxin zearalenone in the Fusarium oxysporum isolate. The Hydrastis root rhizosphere effect provided a selective advantage to the Fusarium isolates closely associated with the root tissue in comparison with the Fusarium isolate that had never been exposed to Hydrastis.     

Increased Terpenoid Accumulation in Cotton (<Emphasis Type="Italic">Gossypium hirsutum</Emphasis>) Foliage is a General Wound Response

The subepidermal pigment glands of cotton accumulate a variety of terpenoid products, including monoterpenes, sesquiterpenes, and terpenoid aldehydes that can act as feeding deterrents against a number of insect herbivore species. We compared the effect of herbivory by Spodoptera littoralis caterpillars, mechanical damage by a fabric pattern wheel, and the application of jasmonic acid on levels of the major representatives of the three structural classes of terpenoids in the leaf foliage of 4-week-old Gossypium hirsutum plants. Terpenoid levels increased successively from control to mechanical damage, herbivory, and jasmonic acid treatments, with E-β-ocimene and heliocide H₁ and H₄ showing the highest increases, up to 15-fold. Herbivory or mechanical damage to older leaves led to terpenoid increases in younger leaves. Leaf-by-leaf analysis of terpenes and gland density revealed that higher levels of terpenoids were achieved by two mechanisms: (1) increased filling of existing glands with terpenoids and (2) the production of additional glands, which were found to be dependent on damage intensity. As the relative response of individual terpenoids did not differ substantially among herbivore, mechanical damage, and jasmonic acid treatments, the induction of terpenoids in cotton foliage appears to represent a non-specific wound response mediated by jasmonic acid. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

NMR Metabolomics of Thrips (<Emphasis Type="Italic">Frankliniella occidentalis</Emphasis>) Resistance in <Emphasis Type="Italic">Senecio</Emphasis> Hybrids

Western flower thrips (Frankliniella occidentalis) has become a key insect pest of agricultural and horticultural crops worldwide. Little is known about host plant resistance to thrips. In this study, we investigated thrips resistance in F ₂ hybrids of Senecio jacobaea and Senecio aquaticus. We identified thrips-resistant hybrids applying three different bioassays. Subsequently, we compared the metabolomic profiles of these hybrids applying nuclear magnetic resonance spectroscopy (NMR). The new developments of NMR facilitate a wide range coverage of the metabolome. This makes NMR especially suitable if there is no a priori knowledge of the compounds related to herbivore resistance and allows a holistic approach analyzing different chemical compounds simultaneously. We show that the metabolomes of thrips-resistant and -susceptible hybrids differed considerably. Thrips-resistant hybrids contained higher amounts of the pyrrolizidine alkaloids (PA), jacobine, and jaconine, especially in younger leaves. Also, a flavanoid, kaempferol glucoside, accumulated in the resistant plants. Both PAs and kaempferol are known for their inhibitory effect on herbivores. In resistant and susceptible F ₂ hybrids, young leaves showed less thrips damage than old leaves. Consistent with the optimal plant defense theory, young leaves contained increased levels of primary metabolites such as sucrose, raffinose, and stachyose, but also accumulated jacaranone as a secondary plant defense compound. Our results prove NMR as a promising tool to identify different metabolites involved in herbivore resistance. It constitutes a significant advance in the study of plant–insect relationships, providing key information on the implementation of herbivore resistance breeding strategies in plants. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Role of Fresh <Emphasis Type="Italic">versus</Emphasis> 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 signals.     

Responses of <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> to Tomato Plants Previously Infected by ToMV or Damaged by <Emphasis Type="Italic">H. armigera</Emphasis>

We report the comparative inducing effects of a phytopathogen and a herbivorous arthropod on the performance of an herbivore. Tomato, Lycopersicon esculentum Mill., was used as the test plant, and tomato mosaic virus (ToMV) and corn earworm, Helicoverpa armigera Hübner, were used as the phytopathogen and herbivore, respectively. There were decreases in the efficiency of conversion of ingested food and efficiency of conversion of digested food when H. armigera was reared on tomato plants that had been previously inoculated with ToMV. However, virus inoculation did not affect feeding or oviposition preferences by H. armigera. In contrast, approximate digestibility, total consumption, relative growth rate, and relative consumption rate were lower for fourth-instar H. armigera that fed on plants previously damaged by the same herbivore. Feeding and oviposition were both deterred for H. armigera that fed on previously damaged plants. The duration of development of H. armigera was also prolonged under this treatment. Infection by ToMV and feeding damage by H. armigera increased the host plant’s peroxidase and polyphenol oxidase activity, respectively, suggesting that the performance of H. armigera may be affected by the induced phytochemistry of the host plant. Overall, this study indicated that, in general, insect damage has a stronger effect than ToMV infection on plant chemistry and, subsequently, on the performance of H. armigera.     

Differential Levels of Insect Herbivory in the Field Associated with Genotypic Variation in Glucosinolates in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Glucosinolates are commonly found in Arabidopsis thaliana and its crucifer relatives, which are known for their role in defense against insect herbivory. In a common garden experiment, we assessed genotypic variation in glucosinolates in A. thaliana and evaluated the association between this chemistry and both plant damage and fitness. Specifically, glucosinolate concentrations were directly associated with damage levels and inversely associated with fitness. These results are contrary to the general expectation that enhanced chemical defense should result in decreased insect herbivory. As the measured insect community in this field trial was dominated by specialist herbivores, this positive relationship between glucosinolates and herbivory agrees with previous observations that glucosinolates (or their hydrolysis products) attract specialist insects. In addition, glucosinolate diversity in this common garden appeared to affect herbivore damage levels. For example, genotypes that contained alkenyl glucosinolates had higher mean damage levels than those that contained hydroxyalkyl glucosinolates. Results suggest that genotypic variation in glucosinolates may be a major factor in determining plant utilization patterns by insect herbivores in the field.     

Spectrum of Cyanide Toxicity and Allocation in <Emphasis Type="Italic">Heliconius erato</Emphasis> and <Emphasis Type="Italic">Passiflora</Emphasis> Host Plants

The larvae of three races of Heliconius erato were fed various species of Passiflora containing varying levels of cyanoglucosides. The mortality rate of larvae and pupae rose when larvae were fed species of Passiflora capable of releasing larger quantities of cyanide. When larvae were fed species of Passiflora with these properties, the resulting adult butterflies also released higher levels of cyanide. This may serve as a defense mechanism. The compounds responsible for the release of cyanide were not evenly distributed throughout the adult butterfly’s body. The thorax contained the highest concentration of cyanogenic substances, followed by the head, wings, and abdomen. The younger tissues of Passiflora plants had higher levels of cyanide-releasing compounds than stems and mature leaves. Cyanogenic glycoside distribution within the plants is consistent with optimal allocation theory. The levels of cyanide-releasing substances in plants varied depending on the season.     

(7<Emphasis Type="Italic">Z</Emphasis>,9<Emphasis Type="Italic">E</Emphasis>)-2-Methyl-7,9-Octadecadiene: A Sex Pheromone Component of <Emphasis Type="Italic">Lymantria Bantaizana</Emphasis>

Our objective was to identify the sex pheromone of Lymantria bantaizana (Lepidoptera: Lymantriidae) whose larvae feed exclusively on walnut, Juglans spp., in China, and Japan. Coupled gas chromatographic–electroantennographic detection (GC-EAD) analyses of pheromone gland extracts revealed a single EAD-active component. Retention index calculations of this compound on four GC columns suggested that it was a methyl-branched octadecadiene with conjugated double bonds. In GC-EAD analyses of 2-methyloctadecenes, (Z)-2-methyl-7-octadecene and (E)-2-methyl-7-octadecene elicited the strongest antennal responses, suggesting that the double bond positions were at C7 and C9. In comparative GC-EAD analyses of pheromone gland extract and stereoselectively synthesized isomers (E,E; E,Z; Z,E; Z,Z) of 2-methyl-7,9-octadecadiene, the (E,Z)- and (Z,E)-isomer had retention times identical to that of the candidate pheromone, but only the latter isomer elicited strong EAD activity. Results of field experiments in Japan substantiated that (7Z,9E)-2-methyl-7,9-octadecadiene is the L. bantaizana sex pheromone, a compound previously unknown in the Lepidoptera. Detection surveys in North America for exotic Eurasian forest defoliators could include traps baited with the L. bantaizana pheromone.     

Relationship Between the Endophyte <Emphasis Type="Italic">Embellisia</Emphasis> spp. and the Toxic Alkaloid Swainsonine in Major Locoweed Species (<Emphasis Type="Italic">Astragalus</Emphasis> and <Emphasis Type="Italic">Oxytropis)</Emphasis>

Locoweeds (Astragalus and Oxytropis spp. that contain the toxic alkaloid swainsonine) cause widespread poisoning of livestock on western rangelands. There are 354 species of Astragalus and 22 species of Oxytropis in the US and Canada. Recently, a fungal endophyte, Embellisia spp., was isolated from Astragalus and Oxytropis spp. and shown to produce swainsonine. We conducted a survey of the major locoweeds from areas where locoweed poisoning has occurred to verify the presence of the endophyte and to relate endophyte infection with swainsonine concentrations. Species found to contain the fungal endophyte and produce substantial amounts of swainsonine were A. wootoni, A. pubentissimus, A. mollissimus, A. lentiginosus, and O. sericea. Astragalus species generally had higher concentrations of swainsonine than Oxytropis. Swainsonine was not detected in A. alpinus, A. cibarius, A. coltonii, A. filipes, or O. campestris. The endophyte could not be cultured from A. mollissimus var. thompsonii or A. amphioxys, but was detected by polymerase chain reaction, and only 30% of these samples contained trace levels of swainsonine. Further research is necessary to determine if the endophyte is able to colonize these and other species of Astragalus and Oxytropis and determine environmental influences on its growth and synthesis of swainsonine.     

Herbivore-Induced Volatiles in the Perennial Shrub, <Emphasis Type="Italic">Vaccinium corymbosum</Emphasis>, and Their Role in Inter-branch Signaling

Herbivore feeding activates plant defenses at the site of damage as well as systemically. Systemic defenses can be induced internally by signals transported via phloem or xylem, or externally transmitted by volatiles emitted from the damaged tissues. We investigated the role of herbivore-induced plant volatiles (HIPVs) in activating a defense response between branches in blueberry plants. Blueberries are perennial shrubs that grow by initiating adventitious shoots from a basal crown, which produce new lateral branches. This type of growth constrains vascular connections between shoots and branches within plants. While we found that leaves within a branch were highly connected, vascular connectivity was limited between branches within shoots and absent between branches from different shoots. Larval feeding by gypsy moth, exogenous methyl jasmonate, and mechanical damage differentially induced volatile emissions in blueberry plants, and there was a positive correlation between amount of insect damage and volatile emission rates. Herbivore damage did not affect systemic defense induction when we isolated systemic branches from external exposure to HIPVs. Thus, internal signals were not capable of triggering systemic defenses among branches. However, exposure of branches to HIPVs from an adjacent branch decreased larval consumption by 70% compared to those exposed to volatiles from undamaged branches. This reduction in leaf consumption did not result in decreased volatile emissions, indicating that leaves became more responsive to herbivory (or “primed”) after being exposed to HIPVs. Chemical profiles of leaves damaged by gypsy moth caterpillars, exposed to HIPVs, or non-damaged controls revealed that HIPV-exposed leaves had greater chemical similarities to damaged leaves than to control leaves. Insect-damaged leaves and young HIPV-exposed leaves had higher amounts of endogenous cis-jasmonic acid compared to undamaged and non-exposed leaves, respectively. Our results show that exposure to HIPVs triggered systemic induction of direct defenses against gypsy moth and primed volatile emissions, which can be an indirect defense. Blueberry plants appear to rely on HIPVs as external signals for inter-branch communication.     

Pollinator and Herbivore Attraction to <Emphasis Type="Italic">Cucurbita</Emphasis> Floral Volatiles

Mutualists and antagonists may place conflicting selection pressures on plant traits. For example, the evolution of floral traits is typically studied in the context of attracting pollinators, but traits may incur fitness costs if they are also attractive to antagonists. Striped cucumber beetles (Acalymma vittatum) feed on cucurbits and are attracted to several volatiles emitted by Cucurbita blossoms. However, the effect of these volatiles on pollinator attraction is unknown. Our goal was to determine whether pollinators were attracted to the same or different floral volatiles as herbivorous cucumber beetles. We tested three volatiles previously found to attract cucumber beetles in a factorial design to determine attraction of squash bees (Peponapis pruinosa), the specialist pollinators of cucurbita species, as well as the specialist herbivore A. vittatum. We found that 1,2,4-trimethoxybenzene was attractive to both the pollinator and the herbivore, indole was attractive only to the herbivore, and (E)-cinnamaldehyde was attractive only to the pollinator. There were no interactions among volatiles on attraction of squash bees or cucumber beetles. Our results suggest that reduced indole emission could benefit plants by reducing herbivore attraction without loss of pollination, and that 1,2,4-trimethoxybenzene might be under conflicting selection pressure from mutualists and antagonists. By examining the attraction of both mutualists and antagonists to Cucurbita floral volatiles, we have demonstrated the potential for some compounds to influence only one type of interaction, while others may affect both interactions and possibly result in tradeoffs. These results shed light on the potential evolution of fragrance in native Cucurbita, and may have consequences for yield in agricultural settings.     

Expression of Constitutive and Inducible Chemical Defenses in Native and Invasive Populations of <Emphasis Type="Italic">Alliaria petiolata</Emphasis>

The Evolution of Increased Competitive Ability (EICA) hypothesis posits that invasive plants in introduced habitats with reduced herbivore pressure will evolve reduced levels of costly resistance traits. In light of this hypothesis, we examined the constitutive and inducible expression of five chemical defense traits in Alliaria petiolata from four invasive North American and seven native European populations. When grown under common conditions, significant variation among populations within continents was found for trypsin inhibitors and peroxidase activity, and glucosinolates and trypsin inhibitors were significantly jasmonate-inducible across populations. Across populations, constitutive levels of glucosinolates and trypsin inhibitors were negatively correlated with their degree of induction, with three North American populations tending to have lower constitutive levels and higher inducibility of glucosinolates than the seven European populations. Alliarinoside and isovitexin 6″-O-β-d-glucopyranoside levels were both higher in North American plants than in European plants, but levels of these compounds were generally increased by jasmonate in European plants and decreased by the same treatment in North American plants. Aside from the tendency for invasive populations to have reduced constitutive glucosinolate levels coupled with increased inducibility, little support for the predictions of EICA was evident in the chemical defenses that we studied.     

Herbivore-Induced Changes in Tomato (<Emphasis Type="Italic">Solanum lycopersicum</Emphasis>) Primary Metabolism: A Whole Plant Perspective

Induced changes in primary metabolism are important plant responses to herbivory, providing energy and metabolic precursors for defense compounds. Metabolic shifts also can lead to reallocation of leaf resources to storage tissues, thus increasing a plant’s tolerance. We characterized whole-plant metabolic responses of tomato (Solanum lycopersicum) 24 h after leaf herbivory by two caterpillars (the generalist Helicoverpa zea and the specialist Manduca sexta) by using GC-MS. We measured 56 primary metabolites across the leaves, stems, roots, and apex, comparing herbivore-attacked plants to undamaged plants and mechanically damaged plants. Induced metabolic change, in terms of magnitude and number of individual concentration changes, was stronger in the apex and root tissues than in undamaged leaflets of damaged leaves, indicating rapid and significant whole-plant responses to damage. Helicoverpa zea altered many more metabolites than M. sexta across most tissues, suggesting an enhanced plant response to H. zea herbivory. Helicoverpa zea herbivory strongly affected concentrations of defense-related metabolites (simple phenolics and precursor amino acids), while M. sexta altered metabolites associated with carbon and nitrogen transport. We conclude that herbivory induces many systemic primary metabolic changes in tomato, and that changes often are specific to a single tissue or type of herbivore. The potential implications of primary metabolic changes are discussed in relation to resistance and tolerance.     

Roquefortine/Oxaline Biosynthesis Pathway Metabolites in <Emphasis Type="Italic">Penicillium</Emphasis> ser. <Emphasis Type="Italic">Corymbifera</Emphasis>: <Emphasis Type="Italic">In Planta</Emphasis> Production and Implications for Competitive Fitness

Three strains of each of the seven taxa comprising the Penicillium series Corymbifera were surveyed by direct injection mass spectrometry (MS) and liquid chromatography–MS for the production of terrestric acid and roquefortine/oxaline biosynthesis pathway metabolites when cultured upon macerated tissue agars prepared from Allium cepa, Zingiber officinale, and Tulipa gesneriana, and on the defined medium Czapek yeast autolysate agar (CYA). A novel solid-phase extraction methodology was applied for the rapid purification of roquefortine metabolites from a complex matrix. Penicillium hordei and P. venetum produced roquefortine D and C, whereas P. hirsutum produced roquefortine D and C and glandicolines A and B. P. albocoremium, P. allii, and P. radicicola carried the pathway through to meleagrin, producing roquefortine D and C, glandicolines A and B, and meleagrin. P. tulipae produced all previously mentioned metabolites yet carried the pathway through to an end product recognized as epi-neoxaline, prompting the proposal of a roquefortine/epi-neoxaline biogenesis pathway. Terrestric acid production was stimulated by all Corymbifera strains on plant-derived media compared to CYA controls. In planta, production of terrestric acid, roquefortine C, glandicolines A and B, meleagrin, epi-neoxaline, and several other species-related secondary metabolites were confirmed from A. cepa bulbs infected with Corymbifera strains. The deposition of roquefortine/oxaline pathway metabolites as an extracellular nitrogen reserve for uptake and metabolism into growing mycelia and the synergistic role of terrestric acid and other Corymbifera secondary metabolites in enhancing the competitive fitness of Corymbifera species in planta are proposed.     

The Desert Locust, <Emphasis Type="Italic">Schistocerca gregaria</Emphasis>, Detoxifies the Glucosinolates of <Emphasis Type="Italic">Schouwia purpurea</Emphasis> by Desulfation

Desert locusts (Schistocerca gregaria) occasionally feed on Schouwia purpurea, a plant that contains tenfold higher levels of glucosinolates than most other Brassicaceae. Whereas this unusually high level of glucosinolates is expected to be toxic and/or deterrent to most insects, locusts feed on the plant with no apparent ill effects. In this paper, we demonstrate that the desert locust, like larvae of the diamondback moth (Plutella xylostella), possesses a glucosinolate sulfatase in the gut that hydrolyzes glucosinolates to their corresponding desulfonated forms. These are no longer susceptible to cleavage by myrosinase, thus eliminating the formation of toxic glucosinolate hydrolysis products. Sulfatase is found throughout the desert locust gut and can catalyze the hydrolysis of all of the glucosinolates present in S. purpurea. The enzyme was detected in all larval stages of locusts as well as in both male and female adults feeding on this plant species. Glucosinolate sulfatase activity is induced tenfold when locusts are fed S. purpurea after being maintained on a glucosinolate-free diet, and activity declines when glucosinolates are removed from the locust diet. A detoxification system that is sensitive to the dietary levels of a plant toxin may minimize the physiological costs of toxin processing, especially for a generalist insect herbivore that encounters large variations in plant defense metabolites while feeding on different species.     

Laboratory Evaluation of <Emphasis Type="Italic">Artemisia annua</Emphasis> L. Extract and Artemisinin Activity against <Emphasis Type="Italic">Epilachna paenulata</Emphasis> and <Emphasis Type="Italic">Spodoptera eridania</Emphasis>

Ethanolic extract of aerial parts of Artemisia annua L. and artemisinin were evaluated as anti-insect products. In a feeding deterrence assay on Epilachna paenulata Germ (Coleoptera: Coccinellidae) larvae, complete feeding rejection was observed at an extract concentration of 1.5 mg/cm² on pumpkin leaf tissue. The same concentration produced a feeding inhibition of 87% in Spodoptera eridania (Cramer) (Lepidoptera: Noctuidae). In a no-choice assay, both species ate less and gained less weight when fed on leaves treated with the extract. Complete mortality in E. paenulata and 50% mortality in S. eridania were observed with extract at 1.5 mg/cm². Artemisinin exhibited a moderate antifeedant effect on E. paenulata and S. eridania at 0.03–0.375 mg/cm². However, a strong effect on survival and body weight was observed when E. paenulata larvae were forced to feed on leaves treated at 0.03 and 0.075 mg/cm². Artemisia annua ethanolic extract of aerial parts at 1.5 mg/cm² showed no phytotoxic effect on pumpkin seedlings.     

Response of <Emphasis Type="Italic">Plutella xylostella</Emphasis> and its Parasitoid <Emphasis Type="Italic">Cotesia plutellae</Emphasis> to Volatile Compounds

The effects of limonene, a mixture of limonene + carvone (1:1, v/v), and methyl jasmonate (MeJA) on diamondback moth (DBM) (Plutella xylostella L.) oviposition, larval feeding, and the behavior of its larval parasitoid Cotesia plutellae (Kurdjumov) with cabbage (Brassica oleracea L. ssp. capitata, cvs. Rinda and Lennox) and broccoli (B. oleracea subsp. Italica cv Lucky) were tested. Limonene showed no deterrent effect on DBM when plants were sprayed with or exposed to limonene, although there was a cultivar difference. A mixture of limonene and carvone released from vermiculite showed a significant repellent effect, reducing the number of eggs laid on the cabbages. MeJA treatment reduced the relative growth rate (RGR) of larvae on cv Lennox leaves. In Y-tube olfactometer tests, C. plutellae preferred the odors of limonene and MeJA to filtered air. In cv Lennox, the parasitoid preferred DBM-damaged plants with limonene to such plants without limonene. C. plutellae females were repelled by the mixture of limonene + carvone. In both cultivars, exogenous MeJA induced the emission of the sesquiterpene (E,E)-α-farnesene, the homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), and green leaf volatile (Z)-3-hexenyl acetate + octanal. The attractive effect of limonene and MeJA predicts that these two compounds can be used in sustainable plant protection strategies in organic farming.