Relationship between deterrence and toxicity of plant secondary compounds for the grasshopperSchistocerca americana

A variety of plant secondary compounds, several of which are quite widespread in nature were tested for their deterrence to the generalist grasshopperSchistocerca americana in short-term behavioral assays. The compounds were coumarin, salicin, tannic acid, gramine, nicotine, quinine, carvone, geraniol, abietic acid, umbelliferone, and ursolic acid. These were then tested for their post-ingestional effects over the whole of the last larval instar. Different methods were employed to mask the taste of compounds that were deterrent in order to ensure that any effects were not due to reduced feeding. In no case was there any indication of a detrimental effect or any trend suggesting one. In two cases, there was a significant increase in growth rate with the addition of the secondary compound to the diet. The evolutionary implications of these findings are discussed.     

Behavioral (Feeding) Responses of the Crayfish, Procambarus clarkii, to Natural Dietary Items and Common Components of Formulated Crustacean Feeds

We have examined the behavioral (feeding) response of Procambarus clarkii to natural dietary items (zooplankton, live fishes, dead fishes, and fish eggs) and common components of formulated feeds used in the aquaculture industry (soybean meal, fish meal, corn meal, alfalfa meal, and vitamin C). The feeding response by P. clarkii was determined using an ordinally ranked, whole-animal bioassay that included the following behaviors: (1) movement of the maxillipeds for longer than three seconds, (2) increased movement of the walking legs with dactyl probing, (3) movement of walking legs to the mouth, and (4) orientation of the entire body towards the odor source. Feeding behavior was determined in response to intact items, bathwater containing aqueous leachates from intact items, water and methanol fractions of bathwater eluted through a C₁₈ resin flash chromatography column, and size fractions of bathwater containing either molecules 10,000 Da or molecules >10,000 Da. All fractions tested were significantly stimulatory. Zooplankton was the most stimulatory of the natural dietary items tested. However, the C₁₈ water fraction of the soybean meal bathwater before size fractionation (containing molecules both <10,000 and >10,000 Da) was the most stimulatory of the common feed components and more stimulatory than the zooplankton. Proximate analysis indicated that the compounds present in this fraction were ca. 51% soluble carbohydrate, 4% soluble protein, and 45% unknown (assumed to be insoluble carbohydrates, insoluble proteins, and ash). We hypothesize that the primary compounds in soybean meal responsible for eliciting a feeding response in P. clarkii are soluble carbohydrates and/or glycoproteins.     

Taste thresholds and suprathreshold responses to tannin-rich plant extracts and quinine in a primate species (Microcebus murinus)

Theories of plant chemical defenses discriminate between quantitative digestibility reducers (e.g., tannins) and qualitative toxins (e.g., alkaloids). Since the differential effect on taste of these compounds is poorly known, we recorded ingestive responses of a primate species, Microcebus murinus, to four tannin-rich plant extracts and to quinine, by using the behavioral method of the two-bottle test. The efficiency of tannic extracts at precipitating protein was measured with the blue BSA method. Inhibition taste thresholds for tannins added to a moderately sweet solution varied between 0.25 and 2 g/l. The threshold for quinine hydrochloride was 0.32 g/l. The profiles of the response/concentration curves established for these astringent and bitter substances were similar, with maximal inhibition of consumption occurring for near-threshold concentrations. The large amounts of quinine required to deter this small-bodied species from feeding were unexpected, given its unspecialized frugivorous/insectivorous diet. We propose that the taste responses of Microcebus to tannins have been shaped in relation to the widespread occurrence of efficient polyphenols in food plants, while low responsiveness to quinine reflects a low risk of ingesting toxic alkaloids when feeding on ripe fruits and insects.     

Microbial decomposition of ferulic acid in soil

The suppression of plant growth by different phenolic acids is well known. This work was designed to determine if ferulic acid, a known phenolic inhibitor of plant growth, accumulates in the soil and if soil microorganisms could be isolated that metabolize it. Over 99% of the extractable ferulic acid was lost from decaying hackberry leaves in 300 days. During this time the amount in the top 15 cm of soil remained fairly constant at about 30 ppm, except for the March sample which was significantly higher than the rest. Addition of ferulic acid to soil caused an increase in CO₂ evolution and in numbers of a select group of microorganisms.Rhodotorula rubra andCepnalosporium curtipes, which actively metabolize ferulic acid, were isolated, but the metabolic pathways employed appear to be different from the reported one. The reported pathway for ferulic acid breakdown is ferulic acid to vanillic acid to protocatechuic acid to -keto-adipic acid.Rhodotorula Rubra was found to convert ferulic acid to vanillic acid, but no evidence was found for utilization of the rest of the pathway.Cephalosporium curtipes appears to use a different pathway or to metabolize intermediate compounds rapidly without accumulating them, because no phenolic compounds were found during the breakdown of ferulic acid. The presence in the soil of microorganisms that metabolize ferulic acid and other phenolic acids is ecologically significant because such organisms prevent long-term accumulations of these substances, which are toxic to many other microorganisms and higher plants.     

Identification of Semiochemicals Released During Aphid Feeding That Attract Parasitoid Aphidius ervi

Herbivore induced release of plant volatiles mediating the foraging behavior of the aphid parasitoid Aphidius ervi was investigated using the pea aphid, Acyrthosiphon pisum, feeding on broad bean, Vicia faba. Behavioral responses were studied using an olfactometer and a wind tunnel. Volatiles obtained by air entrainment of aphid infested plants were more attractive to A. ervi than those from uninfested plants, in both behavioral bioassays. GC-EAG of both extracts showed a number of peaks associated with responses by A. ervi, but with some differences between extracts. Compounds giving these peaks were tentatively identified by GC-MS and confirmed by comparison with authentic samples on GC, using two columns of different polarity. The activity of pure compounds was further investigated by EAG and wind tunnel assays. Results showed that, of the compounds tested, 6-methyl-5-hepten-2-one was the most attractive for A. ervi females, with linalool, (Z)-3-hexen-1-yl acetate, (E)--ocimene, (Z)-3-hexen-1-ol, and (E)--farnesene all eliciting significantly more oriented flight behavior than a solvent control. Foraging experience significantly increased parasitoid responses to these compounds, with the exception of (E)--farnesene. Time-course GC analysis showed that feeding of A. pisum on V. faba induced or increased the release of several compounds. Release of two of these compounds (6-methyl-5-hepten-2-one and geranic acid) was not induced by the nonhost black bean aphid, Aphis fabae. During the analysis period, production of (E)--ocimene remained constant, but 6-methyl-5-hepten-2-one, linalool, geranic acid, and (E)--farnesene appeared during the first day after A. pisum infestation and increased in concentration with increasing time of aphid feeding.     

The role of leaf lipids in food selection by larvae of the tobacco hornworm,Manduca sexta

The role of leaf lipids in food plant selection by larvae ofManduca sexta was investigated by measuring preference responses in a two-choice preference test using glass fiber filter paper disks laced with extract (test) or water (control). The larvae respond to the petroleum ether extract of whole leaves of the host-plantLycopersicon esculentum (tomato) extract in a concentration-dependent manner. At natural concentration it is the most strongly stimulating extract or compound yet tested using the disk test. This response is affected by food plant experience of the larvae, suggesting stimulation by plant-specific compounds in the extract. The extract contains volatile compounds that attract the larvae. In contrast, it does not promote continued feeding on an agar-cellulose diet that incorporates the extract. Also stimulating are the extracts of leaf surfaces of two hosts,L. esculentum andSolanum pseudocapsicum, and two acceptable nonhosts,Brassicae napus andVigna sinensis, indicating the presence of nonpolar feeding stimulants at the leaf surface. However, similar leaf-surface extracts of the unacceptable plantCanna generalis were inactive, although the surface extraction process renders this plant acceptable. Leaf-surface extracts ofL. esculentum, S. pseudocapsicum, andB. napus evoke feeding responses that are qualitatively comparable to those of their corresponding leaves. However, no such parallel is found for surface extracts ofV. sinensis andC. generalis. Thus, nonpolar compounds at the leaf surface of host and some acceptable non-host-plant species strongly stimulate feeding and hence must play an important role in food selection by the tobacco hornworm.     

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.     

Developmental, Gustatory, and Behavioral Responses of Leafroller Larvae, Choristoneura rosaceana, to Tannic Acid and Glucose

Soluble sugars are essential nutrients generally perceived as phagostimulants to most insects studied. However, tannins are known as digestibility reducers, hence deleterious to caterpillar development, and as deterrents as well. Previous work demonstrated that larvae of the polyphagous oblique-banded leafroller, Choristoneura rosaceana, performed better when reared on a control + 0.5% tannic acid diet than on the standard control diet and that larvae reared on a control + 5% glucose diet had slower development and reduced survival. This study was designed to elucidate the behavioral and neurophysiological components of the larval responses to tannic acid and glucose. C. rosaceana larvae were reared individually from the first to the sixth instar on one of four different artificial diets: (1) control; (2) control + 5% glucose; (3) control + 0.5% tannic acid; (4) control + 5% glucose + 0.5% tannic acid. After 14 days, larvae reared on the control + 5% glucose diet had not developed past the fourth instar, whereas a considerable proportion of larvae reared on the control + 0.5% tannic acid diet had already attained the pupal stage. Insects reared on the control or the control + 5% glucose + 0.5% tannic acid diet had intermediate development, with most larvae in the fifth instar. In addition, once the mid-sixth instar was reached, the feeding preferences to 25 and 300 mM glucose, 25 mM tannic acid, and 25 mM glucose + 25 mM tannic acid over water were assessed in two-choice tests. Feeding affected preference. Control-reared insects preferred feeding on treatments containing glucose and were not deterred by tannic acid. However, larvae that had been exposed to tannic acid during their development were deterred by tannic acid and their glucose discrimination was impaired. The sensitivity to glucose was also examined from neurophysiological recordings by stimulating the sugar-sensitive cell (cell 1) on the lateral styloconic sensillum of the maxillary galea with increasing concentrations of glucose (1, 10, 25, 50, 100, 200, 300, and 500 mM). We also determined whether tannic acid was phagostimulatory, since insects develop relatively quickly on a diet containing this compound, by testing 1 mM tannic acid, 1 mM tannic acid + 300 mM glucose, and 300 mM glucose on the lateral styloconic sensilla. The traces indicated that 1 mM tannic acid was not detected by any of the four chemosensory cells in these sensilla. The combination of tannic acid and glucose produced no spikes from the sugar-sensitive cell, whereas a prominent spike activity resulted with 300 mM glucose. We concluded that, although C. rosaceana larvae develop faster on a tannic acid diet, this compound is not a phagostimulant. The converse is true for glucose; i.e., it stimulates the sugar-sensitive cell in the lateral styloconica in a concentration-dependent fashion. Previous dietary experience changes the sensory and behavioral responses of C. rosaceana to glucose. Our findings imply that not all compounds that are phagostimulatory are necessarily beneficial to an insect's fitness. Therefore, developmental studies should be interpreted in conjunction with behavioral and physiological data.     

Branch-cutting behavior by the vole (Microtus pennsylvanicus)

While investigating the preferences of meadow voles (Microtus pennsylvanicus) toward various coniferous species, we noted they often cut the branches of young trees, and then left them lying on the snow for two days or more before eating them completely. We tested whether this behavioral manipulation of a potential food resource resulted in a decrease in the level of secondary metabolites in branches. We found that after being cut for more than three days, the branches of young conifers had significantly less phenolics and condensed tannins. These postmanipulation levels of secondary metabolites were similar to those found in preferred summer food resources. Voles were thus capable of transforming a low-quality resource into one that was more compatible with their feeding patterns and detoxification mechanisms.     

Chemical Defenses Against Fish Predation in Three Brazilian Octocorals: 11β,12β-Epoxypukalide as a Feeding Deterrent in Phyllogorgia dilatata

The feeding-deterrence properties of crude extracts of three Brazilian octocoral species, Neospongodes atlantica Kükenthal (Alcyonacea, Nephtheidae), Plexaurella regia Castro (Gorgonacea, Plexauridae), and Phyllogorgia dilatata Esper (Gorgonacea, Gorgoniidae), were investigated. All the extracts were incorporated into food strips at the concentrations occurring in the living organisms. Crude extract and its ethyl acetate fraction obtained from P. dilatata collected in Armação dos Búzios (Rio de Janeiro State), when incorporated into artificial diets and tested in the habitat of origin, reduced consumption of food strips by fishes, relative to controls. Crude extracts from two octocoral species collected at the National Marine Park of Abrolhos (Bahia State), N. atlantica and P. regia, had no apparent feeding-deterrence properties; in fact, they seemed to stimulate feeding. Bioassay-guided fractionation of the bioactive P. dilatata crude extract revealed that the deterrent property was restricted to a medium polarity fraction. Field palatability experiments with two pure compounds isolated from this fraction revealed that the furanocembranolide 11,12-epoxypukalide is a potent feeding deterrent produced by P. dilatata against fish. Apparently, furanocembranolides are a particular class of compounds with feeding deterrent properties, protecting some octocorals from potential fish predator species in both tropical and temperate environments.     

Ethyl m-Digallate from Red Maple, Acer rubrum L., as the Major Resistance Factor to Forest Tent Caterpillar, Malacosoma disstria Hbn.

An ethanolic extract of red maple (Acer rubrum L.) leaves (RME) applied to trembling aspen (Populus tremuloides Michx.) leaves reduced feeding in choice test assays with forest tent caterpillar larvae (Malacosoma disstria Hbn.) (FTC), whereas a trembling aspen foliage extract, similarly applied, stimulated feeding. Compounds isolated from the RME were gallic acid, methyl gallate, ethyl gallate, m-digallate, ethyl m-digallate, 1-O-galloyl--D-glucose, 1-O-galloyl--L-rhamnose, kaempferol 3-O--D-glucoside, kaempferol 3-O--D-galactoside, kaempferol 3-O--L-rhamnoside, kaempferol-3-O-rhamnoglucoside, quercetin 3-O--D-glucoside, quercetin 3-O--L-rhamnoside and quercetin 3-O-rhamnoglucoside, (–)-epicatechin, (+)-catechin and ellagic acid. All of the gallates, (–)-epicatechin, and kaempferol 3-O--L-rhamnoside deterred feeding on trembling aspen leaf disks when applied at 0.28 mg/cm². The two digallates deterred feeding by 90% and were the most effective. HPLC analysis indicated that ethyl m-digallate is present in amounts 10–100 × higher in RME (2.5–250 mg/g) than any other compound. Thus, ethyl m-digallate appears to be the major compound protecting red maple from feeding by FTC, with a minor contribution from other gallates.     

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.     

Phytotoxicity of Vulpia Residues: II. Separation, Identification, and Quantitation of Allelochemicals from Vulpia myuros

Phytotoxicity-based extraction and fractionation were employed to separate and purify the allelochemicals from an aqueous extract of vulpia (Vulpia myuros) residues. Further analyses, identification, and quantitation of these allelochemicals were conducted by gas chromatography–mass spectrometry on the most toxic fraction. Twenty-one compounds were identified in the ether fraction from the vulpia residue aqueous extract. They were catechol, hydroquinone, 3,4-dimethoxyphenol, pyrogallol, coniferyl alcohol, and benzoic, succinic, hydrocinnamic, salicylic, protocatechuic, vanillic, gentisic, syringic, p-hydroxybenzoic, -hydroxybenzenepropanoic, p-hydroxyphenylacetic, p-hydroxybenzenepropanoic, hydroferulic, p-coumaric, hydrocaffeic, and ferulic acids. A chromatographic internal standard method with multiple-point calibration graphs was used to quantify the identified compounds. Quantities in the vulpia residues ranged from 1.36 to 81.0 g/g dry residue, in total accounting for 0.05% of the dry weight residue. The combined syringic and hydroferulic acids were present in the largest amount, 140.11 g/g residue. Vanillic, succinic, p-hydroxybenzenepropanoic, and salicylic acids were next in amount, ranging from 37.24 to 81.24 g/g residue. Catechol, hydrocinnamic acid, and hydroquinone were present in the smallest quantities, ranging from 1.36 to 1.82 g/g residue. The remainder of the compounds were intermediate in amount, ranging from 2.33 to 18.1 g/g residue.     

Feeding response of Ips paraconfusus to phloem and phloem metabolites of Heterobasidion annosum-inoculated ponderosa pine,Pinus ponderosa

In studies of feeding by the bark beetle, Ips paraconfusus, two pine stilbenes (pinosylvin and pinosylvin methyl ether), ferulic acid glucoside, and enantiomers of the four most common sugars present in ponderosa pine phloem (sucrose, glucose, fructose, and raffinose) did not stimulate or reduce male feeding when assayed on wet -cellulose with or without stimulatory phloem extractives present. When allowed to feed on wet -cellulose containing sequential extracts (hexane, methanol, and water) of ponderosa pine phloem, methanol and water extractives stimulated feeding, but hexane extractives did not. Males confined in wet -cellulose containing aqueous or organic extracts of culture broths derived from phloem tissue and containing the root pathogen, Heterobasidion annosum, ingested less substrate than beetles confined to control preparations. In an assay using logs from uninoculated ponderosa pines, the mean lengths of phloem in the digestive tracts increased as time spent feeding increased. Males confined to the phloem of basal logs cut from ponderosa pines artificially inoculated with H. annosum ingested significantly less phloem than beetles in logs cut from trees that were (combined) mock-inoculated or uninoculated and did not contain the pathogen. However, individual pathogen-containing treatments were not significantly different from uninoculated controls. It was concluded that altered feeding rates are not a major factor which may explain why diseased ponderosa pines are colonized by I. paraconfusus.     

Behavioral Effects and Sensory Detection of Drimane Deterrents in Myzus persicae and Aphis gossypii Nymphs

Eleven synthetic drimane compounds were tested for their deterrency to nymphs of Myzus persicae (Sulzer) and Aphis gossypii (Glover) (Homoptera: Aphididae). In general, A. gossypii nymphs were less sensitive to the drimanes than M. persicae nymphs. Warburganal (1) and polygodial (2), with a -dialdehyde configuration and double bond at C7–C8, were highly active as deterrents and/or feeding inhibitors against both species. One of the synthetic compounds (10) showed relatively high activity against A. gossypii, while it was not active at all against M. persicae, indicating that synthetic analogs of natural deterrents can be highly selective feeding deterrents. From 24-hr interval observations and ablation studies, it is concluded that nymphs of both M. persicae and A. gossypii detect polygodial and possibly the other drimanes tested with contact chemosensilla at the tips of their antennae. The ablation studies also showed that in both species no tarsal, labial, or epipharyngeal sensilla are involved in detecting polygodial within 24 hr.     

(1S,2R,4S,5S)-angelicoidenol-2-o-β-d-glucopyranoside—A moose deterrent compound in Scots pine (Pinus sylvestris L.)

Hydrophilic extracts of Scots pine (Pinus sylvestris L.) twigs have been investigated for palatability to moose in feeding experiments. The predominant repellent effect was observed from the 2-O--d-glucopyranoside of the monoterpene (1S,2R,4S,5S)-angelicoidenol. Of other isolated and tested substances, only taxifolin-3-O--d-glucopyranoside had a deterring effect. One fraction-containing mainly aliphatic compounds-also had a strong effect, but the effect was lost after further fractionation. Tannin-containing fractions had no effect. Angelicoidenol glucoside has not previously been reported fromP. sylvestris. The results are discussed in view of phenols as defense compounds.     

Spring migration of damson-hop aphid,Phorodon humuli (Homoptera,Aphididae), and summer host plant-derived semiochemicals released on feeding

Behavioral studies using an olfactometer demonstrated that spring migrants of the damson-hop aphid,Phorodon humuli, respond to semiochemicals released by spring migrants feeding on hop leaves. Samples of the total volatiles released on feeding were analyzed by coupled gas chromatographysingle cell recording techniques and showed the presence of several active components. Three compounds were identified, using coupled gas chromatography-mass spectrometry, as methyl salicylate, (E)-2-hexenal and-caryophyllene. These three compounds elicited responses from separate olfactory receptors on the antenna. In the olfactometer, both (E)-2-hexenal and-caryophyllene gave positive responses from spring migrants, and a mixture of the two compounds in the natural ratio was more attractive than (E)-2-hexenal alone. Addition of methyl salicylate eliminated the response to the active binary mixture.     

Isolation and Identification of Primary Metabolite Feeding Stimulants for Adult Western Corn Rootworm, Diabrotica virgifera virgifera Leconte, from Host Pollens

Consumption of pollens from several host plants was quantified for adult western corn rootworm, Diabrotica virgifera virgifera, with significantly more sweet corn and winter squash pollen consumed than that of common sunflower and Canada goldenrod. Sequential solvent extraction and chromatographic fractionation of host pollens, guided by a feeding bioassay of respective fractions, isolated dominant phagostimulatory activity in pools of free amino acids. Amino acid profiles were determined by TLC and HPLC analysis. For sweet corn pollen, proline was predominant, accounting for 70% of a total 95.3 mol free amino acid/g pollen; serine, alanine, -aminobutyric acid (GABA), and asparagine occurred in decreasing order of concentration. In squash pollen, -alanine predominated (30% of 111 mol amino acid/g pollen), followed by asparagine, alanine, proline, and GABA. Histidine (42% of 83.8 mol amino acid/g pollen), proline, aspartic acid, asparagine, and alanine were dominant for sunflower pollen; and for goldenrod, proline (51% of 50.6 mol amino acid/g pollen), trans-hydroxyproline, serine, histidine, and alanine predominated. The five dominant amino acids from each pollen accounted for 80–90% of total amino acid content. Simple mixtures of some dominant amino acids, approximating levels found in sweet corn and squash pollen, gave phagostimulation similar to that of the crude extracts.     

Mammalian exocrine secretions XVI. Constitutents of secretion of supplementary sacculi of dwarf hamster, Phodopus sungorus sungorus

As a first step in a study of the role of the secretion of the supplementary sacculi (buccal secretion) of the dwarf hamster, Phodopus sungorus sungorus, almost complete chemical characterization of the secretion was achieved. The 35 compounds identified include carbon dioxide, hydrogen sulfide, a large number of carboxylic acids (representing the bulk of the organic volatile fraction of the secretion), phenol, 2-piperidone, indole, two long-chain hydroxyesters, cholestrol, desmosterol, and lanosterol. The position of the double bonds in -icosadienyl--butyrolactone and -henicosadienyl--butyrolactone could not be determined, and these two compounds remained only partially characterized. Large variations were found in the relative concentrations in which the short-chain carboxylic acids are present in the secretions of individual animals, and although this aspect was not investigated in sufficient detail in the present investigation, the difference in the carboxylic acid profiles of the secretions of individual animals could play a role in individual recognition in this animal.     

The role of phytoecdysteroids in bracken fern,Pteridium aquilinum (L.) Kuhn as a defense against phytophagous insect attack

Analysis of green bracken fronds collected during July, August, and October, 1975, for phytoecdysteroids showed that these compounds occur in only trace amounts (0.25–0.53 g/kg fresh weight [FW]). The effect of ecdysteroids on the feeding behavior of seven species of insect showed that four species were deterred at ecdysteroid concentrations at or above 60 mg/kg FW diet; one species of insect at 6 mg/kg or above, and two species which were not affected at the higher concentrations. It was concluded that the levels of phytoecdysteroids in bracken would not deter insects from feeding on the plant. The previously published data relevant to the possible role of phytoecdysteroids as defense compounds are also discussed.