Influence of foliar glucosinolates in oilseed rape and mustard on feeding and growth of the bertha armyworm,Mamestra configurata Walker

The relationship between host plant glucosinolate profile and feeding and growth of the Bertha armyworm,Mamestra configurata Walker was investigated using eight cultivated rape and mustard varieties. Mean larval weights of neonates reared on intact rosette-stage plants were significantly different on the different species in the orderBrassica juncea <Sinapis alba <B. napus <B. campestris. WhileB. juncea was least preferred,S. alba was significantly more attractive to neonate larvae in choice tests. Relative consumption and growth rates of fourth-instar larvae were also reduced onB. juncea foliage. Other differences were dependent on the plant growth stage. Neonate preference was not correlated to total glucosinolate levels, but rather to the concentrations of isothiocyanate-releasing glucosinolates. However, the relationship between consumption and glucosinolate levels was inconsistent. Relative growth rate was negatively correlated to total glucosinolate content for stage 3 and 4 foliage—mainly due to the concentration of isothiocyanatereleasing glucosinolates. The relative importance of isothiocyanate-releasing glucosinolates was verified by rearing neonates on meridic diets containing equimolar concentrations of sinigrin, its metabolite, allyl isothiocyanate, and indole-3-carbinol, metabolite of 3-indolylmethyl glucosinolate. Sinigrin and allyl isothiocyanate in the diet produced virtually identical negative weight vs. concentration regression lines. No such dose-response effect was observed with indole-3-carbinol. The data suggest that foliar isothiocyanate-releasing glucosinolates may provide some degree of plant protection from polyphagous insects.     

Feeding preferences of spruce budworm (Choristoneura fumiferana Clem.) larvae to some host-plant chemicals

Single-choice and three-choice tests were used to determine the relative importance of host-plant chemical extracts in eliciting feeding by spruce bud worm larvae. Water-soluble components of the host trees are the most important and, of these, the sugar and glycoside fractions are the most stimulating. The amino acid and organic base fractions have no apparent effect. The organic acid fractions deter feeding slightly.Lepidoptera: Tortricidae.Supported by the National Sciences and Engineering Research Council of Canada, Operating Grant No. A9723 to P.J.A.     

Variable diets and changing taste in plant-insect relationships

The host ranges of phytophagous insects are determined to a large degree by plant chemistry. Specialist insects are often closely associated with plants that produce characteristic chemicals, which may act as attractants or stimulants to aid in finding or recognizing a host. Generalist insects are generally believed to rely on the presence of repellents or deterrents to ensure avoidance of unsuitable plants. However, the chemistry of any plant can be highly variable, as a result of growth characteristics, genetic variation, or environmental factors. Such variable chemistry may provide windows of opportunity for nonadapted insects to utilize a plant or for a plant to become resistant to a normally adapted herbivore. Differences in insect responses to plant constituents may also result from genetic variation or environmental factors. In particular, dietary experience has been found to influence the ability of insects to taste plant chemicals that may serve as signals of suitability or unsuitability. Certain dietary constituents appear to suppress the development of taste sensitivity to deterrents in an insect, whereas the presence of specific stimulants in the diet may result in the development of dependence on these compounds. These findings further emphasize the fact that the dynamics of plant biochemistry along with plasticity in the sensory system of insects might be expected to play a major role in the evolution of new plant–insect relationships.     

Developmental profile of sinalbin (p-hydroxybenzyl glucosinolate) in mustard seedlings,Sinapis alba L., and its relationship to insect resistance

Sinalbin was identified as a chemical component of insect anti-xenosis and antibiosis resistance mechanisms in seedlings ofSinapis alba by DEAE-Sephadex chromatography, HPLC, treatment with sulfatase and myrosinase, various feeding tests using artificial and natural substrates, and by measuring sinalbin concentrations in cotyledons and leaves during seedling development. The effects of sinaibin on feeding were dependent upon the insect species and upon the rapidly changing profile of sinaibin concentrations in the developing seedling. The high concentrations of sinalbin found in young cotyledons (up to 20 mM) and leaves (up to 10 mM) deterred the feeding of the flea beetle,Phyllotreta cruciferae Goeze and larvae of the bertha armyworm,Mamestra configurata Walker. The protection that sinalbin confers upon the vulnerable, newly emerged seedling (and upon tiny, young leaves) appears critical for the first few days of survival ofS. alba under feeding pressure from flea beetles in the field. The lower concentrations of sinaibin found in older cotyledons and leaves (2–3 mM) offer little or no protection againstP. cruciferae and may actually stimulate the feeding of this crucifer specialist. These concentrations of sinaibin, however, are still effective in reducing the level of feeding by larvae of the more generalist feederM. configurata.     

Evaluation of Dimboa Analogs as Antifeedants and Antibiotics Towards the Aphid Sitobion avenae in Artificial Diets

A total of 25 compounds including benzoxazinones, benzoxazolinones, and N-glyoxylamide derivatives were tested as antifeedants and antibiotics towards the aphid Sitobion avenae in diet bioassays. The antifeedant and mortality indexes increased with the presence of electron-donating groups in the 7 position of the benzoxazinone moiety, the replacement of the oxygen atom by sulfur in the heterocyclic ring, the presence of a hemiacetal instead of an acetal at C-2 of the benzoxazine moiety (and hence the possibility of ring opening), and the presence of a hydroxyl group at C-4 of the benzoxazine moiety (hydroxamic acid) instead of a hydrogen atom (lactam). The results support earlier hypotheses on the chemical bases for the mode of action of these compounds.     

Effects of Plant Flavonoids on Fecundity, Survival, and Feeding of the Formosan Subterranean Termite

Fecundity, mortality, and food consumption of the Formosan subterranean termite, Coptotermes formosanus Shiraki, were evaluated in response to five plant flavonoids (genistein, biochanin A, apigenin, quercetin, and glyceollin). Apigenin fed at 50 g/primary reproductive pair proved to be the most toxic flavonoid. Biochanin A was most effective in reducing fecundity. Subsequently, these two flavonoids were tested through oral feeding and topical application at 100-g dose. Significant reduction in the numbers of progeny was evident for biochanin A in both treatment methods. Choice feeding tests with termite workers showed that initially termites were attracted to filter paper treated with biochanin A, but over a period of 72 hr, consumed significantly less material when compared to controls. Biochanin A is a promising phytochemical with ability to reduce fecundity in primary reproductives of the Formosan subterranean termite, but it does not elicit phagostimulant activity.     

Using wood extracts to determine the feeding preferences of the western drywood termite,Incisitermes minor (Hagen)

Nymphs of the western drywood termite,Incisitermes minor (Hagen), were tested for their feeding preferences on wood and extracts of 11 tree species. The amount of wood consumed was inversely proportional to its specific gravity, but methanol extracts of least preferred woods were also least preferred when termites were confined to paper treated with extracts. In choice tests, only paper treated with extract of wood from which the colony developed and untreated and methanol-treated controls were significantly fed upon. Ponderosa pine extract applied to Douglas fir and sugar pine significantly decreased the amount of wood consumed. Resistance to termite feeding appeared to depend on the presence of repellent chemicals in the wood.     

Feeding and boring behavior of the bark beetle,Ips paraconfusus,in extracts of ponderosa pine phloem

MaleIps paraconfusus Lanier bored and fed in cellulose powder substrates treated with solvent extracts of ponderosa pine phloem in preference to cellulose powder alone. Stimuli that elicit boring and feeding behavior occurred in the water extracts and the combined solvent extracts. No significant boring or feeding occurred in the methanol extract. There was a preference for, but no significant feeding in, the water partition of the ether extract. Feeding, but no preferential boring, occurred in the ether extracts.Coleoptera: Scolytidae.Research supported by the Rockefeller Foundation, an NIH Training Grant, USDA/SEA Regional Research Project W-110, and the USDA Forest Service.From a thesis submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy, by J.S.E., 1979, University of California, Berkeley, California.     

Evaluation of Synergism in the Feeding Deterrence of Some Furanocoumarins on Spodoptera littoralis

The phagodepression activity of five furanocoumarins (FC), bergapten, xanthotoxin, psoralen, imperatorin, and angelicin, has been studied against larvae of Egyptian cotton leafworm Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) using a leaf-disk choice bioassay. The dose range used was 0–10 g/cm² for linear furocoumarins and 0–30 g/cm² for angelicin, the angular furanocoumarin. Dose–feeding deterrency activity correlations were governed by various sigmoidal functions, except in the case of imperatorin and bergapten, which had dose–response curves showing irregular traces with two maxima. All five FCs induced various degrees of phagodepression in S. littoralis; at 3 g/cm², the relative feeding deterrence was bergapten = xanthotoxin > psoralen = imperatorin = angelicin. Structure–activity correlations indicated that a methoxy group on C-5 or C-8 enhanced the activity. Comparison of experimental feeding deterrence of binary mixtures of imperatorin with xanthotoxin, bergapten, or psoralen at various relative concentrations with the calculated additive activity of each combination indicated that a proportion of 40% or more of imperatorin may exert a greater antifeedant effect on S. littoralis than the sum of individual compounds. The effect of angelicin also was examined in combination with psoralen, both of which are present in Psoralea plants. Their mixtures yielded a clear synergistic effect in the 20–80% angelicin range (coactivity coefficient = 124–133). By contrast, the effect of angelicin on xanthotoxin was only additive. In the dose–response curves of imperatorin/bergapten combinations, synergism was found at >60% imperatorin relative concentration and 1 g/cm², whereas lower proportions led to antagonism. The threshold between the two opposing effects was found to depend on the total FC concentration employed. Some natural systems contain FCs in the range of synergistic proportions recorded here and, thus, may have been produced by the host to increase its defensive effect at a lower metabolic cost.     

Volatile Emissions Triggered by Multiple Herbivore Damage: Beet Armyworm and Whitefly Feeding on Cotton Plants

Plants are commonly attacked by more than one species of herbivore, potentially causing the induction of multiple, and possibly competing, plant defense systems. In the present paper, we determined the interaction between feeding by the phloem feeder silverleaf whitefly (SWF), Bemisia tabaci Gennadius (B-biotype = B. argentifolii Bellows and Perring), and the leaf-chewing beet armyworm (BAW), Spodoptera exigua Hübner, with regard to the induction of volatile compounds from cotton plants. Compared to undamaged control plants, infestation with SWF did not induce volatile emissions or affect the number and density of pigment glands that store volatile and nonvolatile terpenoid compounds, whereas infestation by BAW strongly induced plant volatile emission. When challenged by the two insect herbivores simultaneously, volatile emission was significantly less than for plants infested with only BAW. Our results suggest that tritrophic level interactions between cotton, BAW, and natural enemies of BAW, that are known to be mediated by plant volatile emissions, may be perturbed by simultaneous infestation by SWF. Possible mechanisms by which the presence of whiteflies may attenuate volatile emissions from caterpillar-damaged cotton plants are discussed.     

Orientation and Feeding Responses of the Pollen Beetle, Meligethes aeneus, to Candytuft, Iberis amara

The pollen beetle, Meligethes aeneus, which is an important pest of oilseed rape, Brassica napus, and turnip rape, B. rapa var. campestris, does not oviposit in all species of the Brassicaceae. The relationship between M. aeneus and candytuft, Iberis amara (Brassicacae), was investigated as part of chemical ecological studies into the development of control methods employing non-host-derived repellents. In choice and nonchoice feeding tests, M. aeneus completely rejected I. amara. However, in a field experiment using traps baited with flowering racemes of I. amara and B. napus, M. aeneus was attracted to both species. Gas chromatographic (GC) and GC-electroantennogram (GC-EAG) analyses indicated that the profiles of the floral volatiles of the two species are different. At least 12 compounds among the I. amara floral volatiles were detected by the M. aeneus antenna, and, of these, hexanoic acid, (E)-4,8-dimethyl-1,3,7-nonatriene and alpha-cedrene were not found among B. napus flower volatiles. Since M. aeneus is stimulated by floral volatiles to approach I. amara, but rejects it near, or at, the plant surface, I. amara does not produce repellents that could be used to manipulate M. aeneus. However, it may contain feeding deterrent(s) that could be used in "push-pull" control techniques or in the development of resistant brassicaceous crops.     

Effects ofneo-clerodane diterpenes fromTeucrium on feeding behavior of colorado potato beetle larvae

Treatment of potato leaf disks with fourneo-clerodane diterpenes fromTeucrium (eriocephalin, teucrin-A, teucvin, and teuscorolide) significantly reduced feeding by larvae ofLeptinotarsa decemlineata (Colorado potato beetle). Choice and no-choice tests suggest that teuscorolide acts as a feeding deterrent, whereas the antifeedant activity of teucrin-A, teucvin, and eriocephalin is likely associated with a toxic mode of action. Nutritional tests, antifeedant simulation assays, and posttreatment studies confirmed that teucrin-A can be categorized by its mode of action as a toxin, rather than as a feeding deterrent.     

Effects of secondary metabolites from marine algae on feeding by the sea urchin,Lytechinus variegatus

A bioassay was developed to test the hypothesis that secondary metabolites from marine algae affect feeding by sea urchins. During experiments measuring chemoreception and gustation, feeding by the regular sea urchin,Lytechinus variegatus (Lamarck), was inhibited by extracts from the green marine alga,Caulerpa prolifera (Forsskal) Lamouroux andCymopolia barbata (Linneaus) Lamouroux. Cymopol, a monoterpene-bromohydroquinone component ofC. barbata, was partially responsible for the inhibited feeding observed in tests of theCymopolia crude extract. Caulerpenyne, an oxygenated sesquiterpene fromC. prolifera, was responsible for all of the urchin feeding inhibition observed in tests of theC. prolifera crude extract. Feeding was not affected by: (1) extracts from several otherCaulerpa species includingC. mexicana (Sonder) J. Agardh,C. ashmaedii Harvey,C. racemosa v.macrophysa (Kutzing) Taylor, andC. racemosa v.laetevirens (Montagne) Weber-van Bosse; (2) caulerpin, an indole-containing pigment isolated from all of theCaulerpa species exceptC. mexicana; and (3) an extract from the red marine alga,Gracilaria foliifera v.angustissima (Harvey) Taylor, which has no known secondary metabolites. Feeding inhibition was independent of the test diameter which correlated with the reproductive state of the urchins. Feeding inhibition was also independent of the starvation periods between experiments, and the temperature and salinity in ranges tolerated byL. variegatus obtained from the Florida Gulf Coast. The data strongly suggest that at least one alga,Caulerpa prolifera, is chemically defended against a dominant omnivore in its community.     

<Emphasis Type="Italic">Barbarea vulgaris</Emphasis> Glucosinolate Phenotypes Differentially Affect Performance and Preference of Two Different Species of Lepidopteran Herbivores

The composition of secondary metabolites and the nutritional value of a plant both determine herbivore preference and performance. The genetically determined glucosinolate pattern of Barbarea vulgaris can be dominated by either glucobarbarin (BAR-type) or by gluconasturtiin (NAS-type). Because of the structural differences, these glucosinolates may have different effects on herbivores. We compared the two Barbarea chemotypes with regards to the preference and performance of two lepidopteran herbivores, using Mamestra brassicae as a generalist and Pieris rapae as a specialist. The generalist and specialist herbivores did not prefer either chemotype for oviposition. However, larvae of the generalist M. brassicae preferred to feed and performed best on NAS-type plants. On NAS-type plants, 100% of the M. brassicae larvae survived while growing exponentially, whereas on BAR-type plants, M. brassicae larvae showed little growth and a mortality of 37.5%. In contrast to M. brassicae, the larval preference and performance of the specialist P. rapae was unaffected by plant chemotype. Total levels of glucosinolates, water soluble sugars, and amino acids of B. vulgaris could not explain the poor preference and performance of M. brassicae on BAR-type plants. Our results suggest that difference in glucosinolate chemical structure is responsible for the differential effects of the B. vulgaris chemotypes on the generalist herbivore. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Identification and Function of Prothoracic Exocrine Gland Steroids of the Dytiscid Beetles Graphoderus cinereus and Laccophilus minutus

The chemical composition of the prothoracic defensive secretion of the water beetles Graphoderus cinereus and Laccophilus minutus was examined by gas chromatographic–mass spectrometric analysis of trimethylsilylated gland extracts. The main components are 3,11-dihydroxy-5-pregnan-20-one in G. cinereus and 3,12-dihydroxy-5-pregnan20-one in L. minutus. Fully saturated pregnanes have not been found in the prothoracic defensive glands of dytiscids before. Based on tests with synthetic compounds, their biological role was examined in a feeding assay with the minnow Phoxinus phoxinus. Agar pellets containing either steroid strongly deterred feeding.     

Identification of a Triterpenoid Saponin from a Crucifer, Barbarea vulgaris, as a Feeding Deterrent to the Diamondback Moth, Plutella xylostella

Larvae of the diamondback moth, Plutella xylostella, a crucifer specialist, refuse to feed on a crucifer, Barbarea vulgaris, because of the presence of a feeding deterrent, which is extractable with chloroform. We isolated a feeding deterrent from B. vulgaris leaves, by successive fractionations with silica-gel, ODS, i.e., C₁₈ reversed phase, and Sephadex LH-20 column chromatographies, and ODS-HPLC, guided by a bioassay for feeding deterrent activity. The structure of the compound was determined to be a monodesmosidic triterpenoid saponin, 3-O-[O--D-glucopyranosyl-(14)--D-glucopyranosyl]-hederagenin, based on FAB-MS, ¹H- and ¹³C-NMR spectra, and hydrolysis experiments. When the compound was applied to cabbage leaf disks at greater than 0.18 g/mm², consumption of the disks by third instars was less than 11% of control disks treated with the solvent alone. Furthermore, all first instars died on the disks treated with the same concentrations. Because the concentration of the compound in the fresh leaves of B. vulgaris was comparable to the effective dose in the cabbage leaf disk tested, we conclude that the unacceptability of B. vulgaris to P. xylostella larvae is primarily due to this saponin.     

Chemical stimulants and deterrents regulating acceptance or rejection of crucifers by cabbage butterflies

GravidPieris rapae butterflies oviposit on many, but not all, crucifers. Rejection ofErysimum cheiranthoides andCapsella bursa-pastoris was initially explained by the presence of chemical deterrents in the plants. Analyses and bioassays of plant extracts indicated the absence of oviposition stimulants inC. bursa-pastoris, but similar chemical separation ofE. cheiranthoides extracts revealed the presence of stimulants as well as deterrents. Choice tests illustrate how acceptance or rejection of a plant by an insect may depend on the balance of positive and negative chemical stimuli within the plant.     

Cardenolides fromErysimum cheiranthoides: Feeding deterrents toPieris rapae larvae

Larvae of the cabbage butterfly,Pieris rapae, refuse to feed on the wild mustard,Erysimum cheiranthoides, due to the presence of alcoholextractable deterrents. The active components were extracted inton-BuOH, and this extract was separated into four fractions (I–IV) by reverse-phase HPLC. Fractions III and IV retained the feeding deterrent activity. The activity of fraction III was found to be due to the cardenolide diglycosides 1 and 2, which were previously reported as oviposition deterrents for gravidP. rapae butterflies. Three active compounds were isolated from fraction IV by column chromatography on silica gel followed by reverse-phase HPLC. These compounds were identified as a monoglycoside, digitoxigenin 3-O--D-glucoside (4), and two diglycosides, glucodigigulomethyloside (5) and glucodigifucoside (6). An additional cardenolide isolated from fraction II was identified as cheirotoxin (7). All compounds were identified by UV, NMR (¹H and¹³C), and mass spectrometry, as well as hydrolysis experiments. The feeding deterrent activity of these compounds was compared with that of related commercially available chemicals and other compounds isolated fromE. cheiranthoides.