Selective sequestration of milkweed (Asclepias sp.) cardenolides inOncopeltus fasciatus (Dallas) (Hemiptera: Lygaeidae)

The cardenolide content of the gut, wings, and fat body ofOncopeltus fasciatus was examined. The female fat body contained 4–5% of the total cardenolide content of the insect. The cardenolide content of male fat body, and gut and wings of both sexes was below the detection limit of the cardenolide assay. Thin-layer chromatography was used to determine the cardenolide array of various tissues and secretions ofO. fasciatus reared on seeds of a single species of milkweed (A. Speciosa) and adult extracts and dorsolateral space fluid ofO. fasciatus reared on seeds of two species of milkweed with different cardenolide arrays (A. speciosa andA. syriaca). Our results indicate that cardenolides are not sequestered in the insect simply on the basis of polarity and that metabolism and differential excretion of cardenolides are involved in the sequestration of cardenolides inO. fasciatus. The similarities in the cardenolide profiles ofO. fasciatus reared on different food sources, and tissues ofO. fasciatus reared on a single food source indicates that there is regulation of the cardenolide array inO. Fasciatus.     

Variation in cardenolide content of the lygaeid bugs,Oncopeltus fasciatus andLygaeus kalmii kalmii and of their milkweed hosts (Asclepias spp.) in central California

A colorimetric assay was used to quantify the amount of cardenolides in the lygaeid bugsOncopeltus fasciatus andLygaeus kalmii kalmii and their milkweed host plants (Asclepias spp.) in central California. The cardenolide content of individual insects, determined in microgram equivalents of digitoxin, varied from zero to over 300 g per insect. Sources of variation of cardenolide content in the insects include interspecific and intraspecific differences in the content of the host plant species and also differences in the content of plant organs on which insects were feeding. This last source of variability may explain temporal variation in the cardenolide content of the insects. Adults ofO. fasciatus, which migrate into California in the late spring and early summer, and adults ofL. k. kalmii, which emerge from winter hibernacula in the early spring, contained small to immeasureable amounts of cardenolides. The colonization pattern ofO. fasciatus on species ofAsclepias in north central California suggests that this species does not maximize its opportunities to sequester large quantities of cardenolides from potential hosts. The emetic potential of lygaeids in California to vertebrate predators is briefly discussed.     

Specialist Weevil, Rhyssomatus lineaticollis, Does Not Spatially Avoid Cardenolide Defenses of Common Milkweed by Ovipositing into Pith Tissue

Rhyssomatus lineaticollis is a milkweed specialist whose larvae feed upon pith parenchyma in ramet stems of the common milkweed, Asclepias syriaca. Compared with other specialist insect herbivores on milkweeds, this curculionid beetle is unusual in that it is cryptically colored and does not sequester cardenolides characteristic of milkweed chemical defense. Based upon optimal defense theory, we predicted that pith tissue would be low in defensive compounds and that oviposition into the pith would spatially avoid cardenolides. We rejected this hypothesis because we found that pith tissue has a relatively high cardenolide concentration compared to cortex, epidermis, and leaf tissues. Moreover, we found total plant cardenolide concentration was lower in plants that contained the beetle eggs. Cardenolide concentrations were different among tissues in intact stems without the pith herbivore compared to stems where it was present. Furthermore, the overall polarity of the cardenolides present varied among plant tissues and between plants with and without R. lineaticollis eggs. Although we found lower concentrations of cardenolide in piths where the eggs were present, the cardenolides present in the pith contained more nonpolar forms, indicating that the plant may be responding to herbivory by increasing toxic efficacy of cardenolide defenses while lowering the total concentration. We suggest that preoviposition behavior by female beetles, which includes feeding on new leaves of the plant, is a mechanism by which females manipulate plant chemistry and assess quantitative and qualitative changes in cardenolide chemistry in response to herbivory prior to oviposition.     

Distribution of autogenous and host-derived chemical defenses inOreina leaf beetles (Coleoptera: Chrysomelidae)

The pronotal and elytral defensive secretions of 10Oreina species were analyzed. Species feeding on Apiaceae, i.e.,O. frigida andO. viridis, or on Cardueae (Asteraceae), i.e.,O. bidentata, O. coerulea, andO. virgulata, produce species-specific complex mixtures of autogenous cardenolides.O. melanocephala, which feeds onDoronicum clusii (Senecioneae, Asteraceae), devoid of pyrrolizidine alkaloids (PAs) in its leaves, secretes, at best, traces of cardenolides. Sequestration of host-plant PAs was observed in all the other species when feeding on Senecioneae containing these alkaloids in their leaves.O. cacaliae is the only species that secretes host-derived PA N-oxides and no autogenous cardenolides. Differences were observed in the secretions of specimens collected in various localities, because of local differences in the vegetation. The other species, such asO. elongata, O. intricata, andO. speciosissima, have a mixed defensive strategy and are able both to synthesize de novo cardenolides and to sequester plant PA N-oxides. This allows a great flexibility in defense, especially inO. elongata andO. speciosissima, which feed on both PA and non-PA plants. Populations of these species were found exclusively producing cardenolides, or exclusively sequestering PA N-oxides, or still doing both, depending on the local availability of food-plants. Differences were observed between species in their ability to sequester different plant PA N-oxides and to transform them. Therefore sympatric species demonstrate differences in the composition of their host-derived secretions, also resulting from differences in host-plant preference. Finally, within-population individual differences were observed because of local plant heterogeneity in PAs. To some extent these intrapopulation variations in chemical defense are tempered by mixing diet and by the long-term storage of PA N-oxides in the insect body that are used to refill the defensive glands.     

Chemical constituents ofErysimum cheiranthoides deterring oviposition by the cabbage butterfly,Pieris rapae

Avoidance ofErysimum cheiranthoides for oviposition byPieris rapae has been attributed to the presence of water-soluble deterrents. The active material was extracted inton-butanol and isolated by a series of HPLC separations. TLC of the active fraction and visualization of individual constituents with Kedde's reagent indicated that cardenolides are responsible for deterring oviposition. UV spectra were also characteristic of cardenolides. Bioassays of selected known cardenolides revealed a general lack of activity, except for cymarin, which was as strongly deterrent as the most prominent cardenolide isolated in pure form fromE. cheiranthoides. The results suggest that cardenolides in this plant can explain its escape from cabbage butterflies, but specific structural features of the glycosides are necessary for oviposition-deterring activity.     

Production of cardiac glycosides by chrysomelid beetles and larvae

Cardenolides were looked for in 17 chrysomelid beetles belonging to 11 genera from three subfamilies, and they were found only inChrysolina andChrysochloa species (Chrysomelinae, Chrysolinini). The food plants of these insects are not known to produce cardenolides. TheChrysochloa and mostChrysolina species secrete a complex mixture of cardenolides, butChrysolina didymata secretes a single compound, andChrysolina carnifex, none. Several quantitative and perhaps qualitative differences were observed in the patterns of cardenolides produced by far distant populations of bothChrysolina polita andC. herbacea, collected in either France and Belgium, or Greece. These differences remain constant from one generation to the other, whatever the food plant is, and appear to be genetic. InC. polita from Greece, the pattern is unchanged after four generations bred in the laboratory onMentha ×villosa, which is known to be without cardenolides. In adults, the cardenolides are released with the secretion of the pronotal and elytral defensive glands, but in the larvae which lack the defensive glands, cardenolides are also produced. The total amount of cardenolides and the complexity of their mixture increases through the life cycle of the insects. The six main cardenolides secreted byC. coerulans were identified as: sarmentogenin, periplogenin, bipindogenin, and their corresponding xylosides.C. didymata secretes only sarmentogenin.     

Microsomal oxidation of allelochemicals in generalist (Spodoptera frugiperda) and semispecialist (Anticarsia gemmatalis) insect

Midgut microsomes prepared from larvae of the fall armyworm [Spodoptera frugiperda (J.E Smith)], a generalist insect, and the velvetbean caterpillar (Anticarsia gemmatalis Hubner), a semispecialist, were used to study their oxidative activity toward a variety of allelochemicals. Allelochemicals such as terpenoids, alkaloids, indoles, glucosinolates, flavonoids, coumarins, cardenolides, phenylpropenes, and a ketohydrocarbon were all metabolized by the microsomal cytochrome P-450 monooxygenases in both species. Fall armyworm microsomes oxidized monoterpenes more favorably than other types of terpenes, indicating a preference for these compounds. In all instances, the oxidative metabolism of these allelochemicals can be induced by dietary allelochemicals such as indole 3-carbinol, indole 3-acetonitrile, menthol, flavone, or peppermint oil ranging from 1.3- to 9.5-fold. In the case of certain triterpenes, tetraterpene, alkaloid, coumarin, and cardenolides, metabolic activity can only be observed after induction. The monooxygenase activities toward these allelochemicals were generally higher in the generalist than in the semispecialist. These findings provide strong evidence that microsomal monooxygenases play an important role in the detoxification of plant toxins and hence host-plant selections in herbivorous insects.     

Sequestration, Maintenance, and Tissue Distribution of Pyrrolizidine Alkaloid N-Oxides in Larvae of Two Oreina Species

Oreina cacaliae and O. speciosissima are leaf beetles that, as larvae and adults, sequester pyrrolizidine alkaloid N-oxides (PAs) as defensive compounds from their host plants Adenostyles alliariae and Senecio nemorensis. As in most Oreina species, O. speciosissima is also defended by autogenously produced cardenolides (mixed defensive strategy), whereas O. cacaliae does not synthesize cardenolides and is exclusively dependent on host-plant-acquired PAs (host-derived defense). Adults of the two Oreina species were found to have the same PA storage capacity. The larvae, however, differ; larvae of O. speciosissima possess a significantly lower capability to store PAs than O. cacaliae. The ability of Oreina larvae to sequester PAs was studied by using tracer techniques with ¹⁴C-labeled senecionine N-oxide. Larvae of the two species efficiently take up [¹⁴C]senecionine N-oxide from their food plants and store the alkaloid as N-oxide. In O. cacaliae, there is a slow but continuous loss of labeled senecionine N-oxide. This effect may reflect the equilibrium between continuous PA uptake and excretion, resulting in a time-dependent tracer dilution. No noticeable loss of labeled alkaloid is associated with molting. Senecionine N-oxide is detectable in all tissues. The hemolymph is, with ca. 50–60% of total PAs, the major storage compartment, followed by the integument, with ca 30%. The alkaloid concentration in the hemolymph is approximately sixfold higher than in the solid tissues. The selectivity of PA sequestration in larvae is comparable to PA sequestration in the bodies of adult beetles.     

Plant-determined variation in the cardenolide content,thin-layer chromatography profiles,and emetic potency of monarch butterflies,Danaus plexippus L. Reared on milkweed plants in California: 2.Asclepias speciosa

The pattern of variation in gross cardenolide concentration of 111Asclepias speciosa plants collected in six different areas of California is a positively skewed distribution which ranges from 19 to 344 g of cardenolide per 0.1 g dry weight with a mean of 90 g per 0.1 g. Butterflies reared individually on these plants in their native habitats ranged from 41 to 547 g of cardenolide per 0.1 g dry weight with a mean of 179 g. Total cardenolide per butterfly ranged from 54 to 1279 g with a mean of 319 g. Differences in concentrations and total cardenolide contents in the butterflies from the six geographic areas appeared minor, and there were no differences between the males and the females, although the males did weigh significantly more than females. The uptake of cardenolide by the butterflies was found to be a logarithmic function of the plant concentration. This results in regulation: larvae which feed on low-concentration plants produce butterflies with increased cardenolide concentrations relative to those of the plants, and those which feed on high-concentration plants produce butterflies with decreased concentrations. No evidence was adduced that high concentrations of cardenolides in the plants affected the fitness of the butterflies. The mean emetic potencies of the powdered plant and butterfly material were 5.62 and 5.25 blue jay emetic dose fifty units per milligram of cardenolide and the number of ED₅₀ units per butterfly ranged from 0.28 to 6.7 with a mean of 1.67. Monarchs reared onA. speciosa, on average, are only about one tenth as emetic as those reared onA. eriocarpa. UnlikeA. eriocarpa which is limited to California,A. speciosa ranges from California to the Great Plains and is replaced eastwards byA. syriaca L. These two latter milkweed species appear to have a similar array of chemically identical cardenolides, and therefore both must produce butterflies of relatively low emetic potency to birds, with important ecological implications. About 80% of the lower emetic potency of monarchs reared on A. speciosa compared to those reared onA. eriocarpa appears attributable to the higher polarity of the cardenolides inA. speciosa. Thin-layer Chromatographie separation of the cardenolides in two different solvent systems showed that there are 23 cardenolides in theA. speciosa plants of which 20 are stored by the butterflies. There were no differences in the cardenolide spot patterns due either to geographic origin or the sex of the butterflies. As when reared onA. eriocarpa, the butterflies did not store the plant cardenolides withR _f values greater than digitoxigenin. However, metabolic transformation of the cardenolides by the larvae appeared minor in comparison to when they were reared onA. eriocarpa. AlthoughA. eriocarpa andA. speciosa contain similar numbers of cardenolides and both contain desglucosyrioside, the cardenolides ofA. speciosa overall are more polar. ThusA. speciosa has no or only small amounts of the nonpolar labriformin and labriformidin, whereas both occur in high concentrations inA. eriocarpa. A. speciosa plants and butterflies also contain uzarigen, syriogenin, and possibly other polar cardenolides withR _f values lower than digitoxin. The cardenolide concentration in the leaves is not only considerably less than inA. eriocarpa, but the latex has little to immeasurable cardenolide, whereas that ofA. eriocarpa has very high concentrations of several cardenolides. Quantitative analysis ofR _f values of the cardenolide spots, their intensities, and their probabilities of occurrence in the chloroform-methanol-formamide TLC system produced a cardenolide fingerprint pattern very different from that previously established for monarchs reared onA. eriocarpa. This dispels recently published skepticism about the predictibility of chemical fingerprints based upon ingested secondary plant chemicals.Lepidoptera: Danaidae.Apocynales: Asclepiadaceae.This study was supported by U.S. National Science Foundation grants DEB 75-14265 and 78-10658 to Amherst College and BSR-8119382 to the University of Florida with L.P. Brower as Principal Investigator and DEB75-14266, DEB78-15419, and DEB-81-19391 to the University of California at Davis with J.N. Seiber as Principal Investigator.     

Seasonal and intraplant variation of cardenolide content in the California milkweed,Asclepias eriocarpa,and implications for plant defense

Root, stem, leaf, and latex samples ofAsclepias eriocarpa collected from three plots in one population at 12 monthly intervals were assayed for total cardenolide content by spectroassay and for individual cardenolides by thin-layer chromatography. From May to September mean milligram equivalents of digitoxin per gram of dried plant were: latices, 56.8 stems, 6.12 > leaves, 4.0 > roots, 2.5. With the exception of the roots, significant changes in gross cardenolide content occurred for each sample type with time of collection during the growing season, whereas variation within this population was found to be small. Labriformin, a nitrogen-containing cardenolide of low polarity, predominated in the latices. Leaf samples contained labriformin, labriformidin, desglucosyrioside, and other unidentified cardenolides. In addition to most of the same cardenolides as the leaves, the stems also contained uzarigenin. The roots contained desglucosyrioside and several polar cardenolides. The results are compared with those for other cardenolide-containing plants, and discussed in relation to anti-herbivore defense based on plant cardenolide content. Arguments are advanced for a central role of the latex in cardenolide storage and deployment which maximizes the defensive qualities of the cardenolides while preventing toxicity to the plant.Research supported by National Science Foundation Grants DEB 75-14266 and DEB 78-15419 (U.C. Davis) and DEB 75-14265, DEB 78-10658, and DEB 80-40388 (U.F.)     

Palatability of aposematic queen butterflies (Danaus gilippus) feeding onSarcostemma clausum (Asclepiadaceae) in Florida

Queen butterflies (Danaus gilippus) are generally considered unpalatable to predators because they sequester and store toxic cardenolides from their larval food plants. However, a major queen food plant in Florida, the asclepiadaceous vineSarcostemma clausum, is shown here to be a very poor cardenolide source, and queens reared on this plant contain no detectable cardenolide. A direct evaluation of queen palatability using red-winged blackbirds indicates thatS. clausum-reared butterflies are essentially palatable to these predators (85% of abdomens entirely eaten), indicating little protection from either cardenolides, other sequestered phytochemicals, or de novo defensive compounds. Wild-caught queens that presumably fed as larvae uponS. clausum and also had access to adult-obtained chemicals, such as pyrrolizidine alkaloids (PAs), were relatively palatable as well (77% of abdomens eaten); they did not differ significantly in palatability from the labreared butterflies. Together, these findings suggest that; (1)S. clausumfed queens are poorly defended against some avian predators, and (2) for the particular queen sample examined, PAs do not contribute substantially to unpalatability. The discovery thatS. clausum-feeding queens are essentially palatable is of additional significance because it compels a reassessment of the classic mimicry relationship between queen and viceroy butterflies. Viceroys have been shown recently to be moderately unpalatable; therefore, the established roles of model and mimic may be reversed in some cases.     

Chemical constituents of an unacceptable crucifer,Erysimum cheiranthoides,deter feeding byPieris rapae

The wild cruciferErysimum cheiranthoides was found to contain extractable constituents that deterred feeding by larvae of the crucifer specialistPieris rapae when applied to cabbage leaf disks in both choice and nochoice bioassays. High-performance liquid chromatography was used to separate the extract into several fractions, two of which retained the feeding deterrent activity of the extract. UV-absorption spectra of the fractions suggested that one contained cardenolides similar or identical to those reported to deter oviposition byP. rapae onE. cheiranthoides. The other active fraction evidently contains a compound that deters larval feeding but not adult oviposition. The results suggest that the chemical defense ofE. cheiranthoides depends on two types of compounds acting on separate developmental stages of the insect.     

Cardenolide sequestration by the dogbane tiger moth (Cycnia tenera; Arctiidae)

Cycnia tenera adults, reared as larvae onAsclepias humistrata, had 10 times higher cardenolide concentrations, and contained 15 times more total cardenolide, than did moths reared onA. tuberosa. Thin-layer chromatography confirmed that each individual cardenolide visualized in the adult moths reared on the former host plant corresponds to one present in the plant, thus demonstrating that the insects' cardenolides are indeed derived from the larval food. Adult weights were significantly greater when the larvae had been fed upon the higher cardenolide plant species,A. humistrata. Similar results for other milkweed-feeding insects have been interpreted by some authors as evidence against a metabolic cost of handling cardenolides. However, such interpretations confound cardenolide differences among milkweed species with other differences in plant primary and secondary chemistry that affect insect growth and development. While the cooccurrence inC. tenera of other noxious chemicals (e.g., alkaloids) is not precluded, cardenolides sequestered from larval host plants have probably contributed to the evolution of visual and auditory aposematism in this species. As the eggs are laid in large clutches and larvae are gregarious, such aposematism may have evolved via kin selection.     

Cardenolide fingerprint of monarch butterflies reared on common milkweed,Asclepias syriaca L.

Monarch butterfly,Danaus plexippus (L.), larvae were collected during August 1983 from the common milkweed,Asclepias syriaca L., across its extensive North American range from North Dakota, east to Vermont, and south to Virginia. This confirms that the late summer distribution of breeding monarchs in eastern North America coincides with the range of this extremely abundant milkweed resource. Plant cardenolide concentrations, assayed by spectrophotometry in 158 samples from 27 collection sites, were biased towards plants with low cardenolide, and ranged from 4 to 229 g/ 0.1 g dry weight, with a mean of 50 g/0.1 g. Monarch larvae reared on these plants stored cardenolides logarithmically, and produced 158 adults with a normally distributed concentration range from 0 to 792 g/0. l g dry butterfly, with a mean of 234 g/0.1 g. Thus butterflies increased the mean plant cardenolide concentration by 4.7. The eastern plants and their resultant butterflies had higher cardenolide concentrations than those from the west, and in some areas monarchs sequestered more cardenolide from equivalent plants. Plants growing in small patches had higher cardenolide concentrations than those in larger patches, but this did not influence butterfly concentration. However, younger plants and those at habitat edges had higher cardenolide concentrations than either older, shaded, or open habitat plants, and this did influence butterfly storage. There were no apparent topographical differences reflected in the cardenolides of plants and butterflies. Twenty-eight cardenolides were recognized by thin-layer chromatography, with 27 in plants and 21 in butterflies. Butterflies stored cardenolides within the more polar 46% of the plantR _d range, these being sequestered in higher relative concentrations than they occurred in the plants. By comparison with published TLC cardenolide mobilities, spots 3, 4, 9, 16, 24 or 25, 26, and 27, may be the cardenolides syrioside, uzarin, syriobioside, syriogenin, uzarigenin, labriformidin, and labriformin, respectively. Cochromatography with cardenolide standards indicated that desglucosyrioside did not occur in the plants but did occur in 70% of the butterflies, and aspecioside was in 99% of the plants and 100% of the butterflies. The polar aspecioside was the single most concentrated and diagnostic cardenolide in both plants and butterflies. ButterflyR _d values were dependent on those of the plant, and both showed remarkable uniformity over the range of areas sampled. Thus contrary to previous reports,A. syriaca has a biogeographically consistent cardenolide fingerprint pattern. The ecological implications of this for understanding the monarch's annual migration cycle are significant.     

Sequestration of distasteful compounds by some pharmacophagous insects

Several pharmacophagous insects have been shown to sequester specific kairomonal substances or their derivatives in their body tissues. Turnip sawflies,Athalia rosae, visit a plant,Clerodendron trichototmum (Verbenaceae), and feed voraciously on the leaf surface. Clerodendrins were characterized as the potent phagostimulants forA. rosae adults. The insect sequesters some of the analogs and becomes extremely bitter on its body surface. Some chrysomelid leaf beetles associated with cucurbitacins were found to store high concentrations of these bitter principles in their body. South American polyphagous beetles,Diabrotica speciosa andCerotoma arcuata, are strongly arrested by root components from the cucurbit plant,Ceratosanthes hilariana, and selectively accumulate 23,24-dihydrocucurbitacin D, effectively gaining bitterness. Similarly, four species of Asian pumpkin leaf beetles belonging to the genusAulacophora were shown to sequester the same compound in body tissue as the major bitter principle. Three phenylpropanoids closely related to methyl eugenol were found to accumulate in the rectal glands of the male Oriental fruit fly,Dacus dorsalis. One of the rectal gland components, 2-allyl-4,5-dimethoxyphenol was shown to be released in the air during courtship. In all of these cases, selectively sequestered compounds strongly deterred feeding by some predators, thus serving as allomones in this context. Kairomonal and pheromonal functions linked with allomonal sequestration by pharmacophagous feeding has also been suggested.     

Identification of food volatiles attractive toGlischrochilus quadrisignatus andGlischrochilus fasciatus (Coleoptera: Nitidulidae)

Seven volatile compounds identified from the headspace of whole wheat bread dough were investigated for their role in attractingGlischrochilus quadrisignatus andG. fasciatus in the field. Traps baited with either whole wheat bread dough or a synthetic seven-component bread dough odor caught similar numbers of these beetles, suggesting that the seven-compound combination could simulate the behavioral effect of bread dough. A series of trials using traps baited with various combinations of these chemicals showed that five compounds were significantly active in attractingG. quadrisignatus andG. fasciatus, but not all were essential for maximum response. The simplest blend eliciting a level of response comparable to the seven-component combination included ethyl acetate, acetaldehyde, ethanol, and racemic 2-methylbutanol, of which ethyl acetate, ethanol, and acetaldehyde were essential and 2-methylbutanol was replaceable with 2-methylpropanol forG. quadrisignatus attraction. Ethyl acetate and ethanol were essential for comparable attraction ofG. fasciatus. The chemical mediation of food finding in G.quadrisignatus andG. fasciatus is discussed in the context of volatile blends characterized for other nitidulid species.     

A new method for fast isolation of insect antifeedant compounds from complex mixtures

Bioassay-guided isolation of bioactive natural substances requires monitoring of all fractionation and purification steps using a bioassay system. This is often a long and tedious process, especially with insect feeding bioassays. We report here a new method, based on the principle of bioautography, for a quick isolation of insect antifeedant compounds. TLC plates, after development, are coated with a thin layer of artificial diet and fed toSpodoptera litura larvae. The location of uneaten areas is then compared with theR _fvalues of the TLC spots, in order to determine rapidly the active fractions. This methods allows for a very fast determination of the most active antifeedant compounds in a complex mixture and considerably speeds up the isolation process. This new method was successfully applied in the study of antifeedant activity of several plant samples, and results are presented here for a model plant,Skimmia japonica (Rutaceae). Using this new method, the compounds responsible for the feeding-deterrent activity, three furanocoumarins (bergapten, xanthotoxin, and oxypeucedanin), were quickly and efficiently identified.     

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.     

Patterns of Bioactivity and Herbivory on Nothofagus Species from Chile and New Zealand

Nothofagus species from Chile and New Zealand were surveyed in the field for invertebrate abundance and leaf feeding damage and in the laboratory for antifeedant activity against leafrollers (Ctenopsteustis obliquana, Epiphyas postvittana), deterrent activity against pea aphid (Acyrthosiphon pisum), insect growth regulatory activity (Oncopeltus fasciatus), nematicidal activity (Caenorhabditis elegans), antibiotic activity (Pseudomonas solanaciarium), and general toxicity. A data matrix indicated that N. alessandri and N. pumilio most likely have a chemical barrier to insect attack as leaves showed low faunal abundance, low herbivory, and activity in the leafroller antifeedant, aphid deterrent, and nematicidal assays. A chemical examination of N. alessandri that used the leafroller antifeedant test to guide the separation yielded an active fraction containing the flavonoid, galangin, and the stilbene, pinosylvin, which appear to act in concert to deter leafroller feeding. The discovery of the phytoalexin, pinosylvin, in the leaves, raises the possibility that Nothofagus in general, and N. alessandri in particular, may have induced chemical defense mechanisms.