Effect of DIMBOA,an aphid resistance factor in wheat,on the aphid predatorEriopis connexa Germar (Coleoptera: Coccinellidae)

DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one), a secondary metabolite found in cereal extracts, confers resistance in wheat to aphids. Its effect on beneficial organisms was tested on larvae of the aphid predatorEriopis connexa Germar. Larvae were fed until pupation on artificial diets to which different concentrations of DIMBOA (2–200g/g diet) were added, as well as on aphids that had been feeding on wheat seedlings with different DIMBOA levels (140–440 g/g fresh tissue). In diets, the effect of DIMBOA was greatest on survival of third-instar larvae and on the duration of the second and fourth instars. When aphids were provided as food, those that had fed on a wheat cultivar with an intermediate DIMBOA level led to a significantly longer larval duration in the predator than did those that fed on either low or high DIMBOA cultivars. Shortest predator development times were obtained with aphid prey that had fed on high DIMBOA seedlings. Higher DIMBOA levels in the plant appear to reduce aphid feeding rates (and rates of DIMBOA ingestion), decreasing aphid survival and minimizing the effect of the toxin on the predator.On leave from Departamento de Entomologia, Universidad Nacional Agraria La Molina, Apartado aéreo 456, Lima, Perú.     

Hydroxamic acid glucosides in honeydew of aphids feeding on wheat

DIMBOA glucoside (2-O-/gb-D-glucopyranosyl-4-hydroxy-7-meth-oxy-1,4-benzoxazin-3-one), the main hydroxamic acid (Hx) in intact wheat plants, was detected in the honey dew ofRhopalosiphum padi feeding on seedlings of six wheat cultivars that differed in their concentration of Hx, suggesting that the chemical circulates in the phloem. Neither the aglucone (DIMBOA) nor its main breakdown product were found in any of the honeydew samples. Honey dew production by aphids caged on seedlings of the wheat cultivars and DIMBOA glucoside concentrations in the honeydew followed biphasic curves when plotted against Hx concentration, suggesting passive ingestion of the chemical from the phloem at low Hx concentrations and limited ingestion due to feeding deterrency by Hx in mesophyll cells at high Hx concentrations. The presence of plant toxins such as Hx glucosides in the phloem sap, the main ingesta of aphids, and in the mesophyll cells, has major implications for plant defense, through a feeding deterrent effect during stylet penetration, and deterrency (antixenosis) along with antibiosis during feeding.     

Resistance of cereals to aphids: The interaction between hydroxamic acids and UDP-glucose transferases in the aphidSitobion avenue (Homoptera: Aphididae)

UPD-glucose transferases are found in the cytosolic and microsomal fractions of the grain aphidSitobion avenae F. Gel filtration and SDSPAGE revealed that the microsomal fraction contained several forms of the enzyme. The molecular weights of the three most active fractions might be 68,000, 66,000, and 36,500. There was a negative correlation between the enzymes' activity in extracts of aphids and the concentration of DIMBOAaglucone in the winter wheat variety fed on by the aphid. A strong inhibition of the activity of the UPD-glucose transferases was observedin vitro at a concentration of DIMBOA as low as 0.01 mM. There was a greater activity of the enzymes in aphids fed on seedlings of susceptible than on moderately resistant wheat cultivars. Prolonged feeding on resistant cultivars resulted in a further reduction in the activity of the aphid's enzymes. The significance for cereal aphids of the role of their UDP-glucose tranferases in the detoxification of plant allelochemicals and adaptation to resistant varieties of cereals is discussed.     

Honeydew analysis for detecting phloem transport of plant natural products

Analysis of honeydew excreted by various sap-sucking insects indicated the presence of certain plant secondary constituents in the phloem of their host plants. Honeydew excreted by mealybugs (Pseudococcus longispinus), living onCastanospermum australe, contained the indolizidine alkaloid castanospermine, a potent -glucosidase inhibitor. Similarly, honeydew from green peach aphids (Myzus persicae), feeding onSenecio vulgaris flower buds, contained the pyrrolizidine alkaloid senecionine, itsN-oxide, and hydrolytic products including retronecine. Cardenolides were detected in the honeydew of oleander aphids (Aphis nerii) feeding on oleander (Nerium oleander), indicating that these compounds are translocated in the phloem. On the other hand, honeydew from greenbugs (Schizaphis graminum), feeding on barley, lacked gramine or related indole metabolites. Similarly MBOA, the breakdown product of DIMBOA, was not detected in the honeydew of greenbugs living on DIMBOA-containing wheat.     

DIMBOA Glucoside, a Wheat Chemical Defense, Affects Host Acceptance and Suitability of Sitobion avenae to the Cereal Aphid Parasitoid Aphidius rhopalosiphi

The influence of hydroxamic acids (Hx), plant secondary metabolites associated with aphid resistance in wheat, on the host acceptance and suitability of the aphid Sitobion avenae to the cereal aphid parasitoid Aphidius rhopalosiphi was evaluated. Aphids showed a reduction in mean relative growth rate and in body size in the wheat cultivar with higher Hx level. Reduction in aphid size was related to a decreased success in avoiding parasitoid oviposition. A minor increase in A. rhopalosiphi developmental time was observed in aphids feeding on the higher Hx cultivar. Experiments with different concentrations of DIMBOA glucoside, the main Hx in wheat, in artificial diets showed an increase in parasitoid developmental time at the highest concentration, with no change in other performance variables. The evidence is discussed in relation to the compatible utilization of host-plant resistance and biological control in integrated pest management.     

Behavioral and electrophysiological responses of natural enemies to synomones from tea shoots and kairomones from tea aphids,Toxoptera aurantii

Olfactometer bioassays and electrophysiological studies showed that the lacewing, Chrysopa sinica, the aphid parasitoid, Aphidius sp., and the coccinellid, Coccinella septempunctata, all responded to volatiles from tea aphids, Toxoptera aurantii, to hexane or ether rinses of tea aphid cuticles, and to synomones released by aphid-damaged tea shoots, as well as to the tea shoot–aphid complex. Each natural enemy spent more time searching on a filter paper treated with tea aphid honeydew than on a blank control filter paper. The interaction between synomones from aphid-damaged shoots and kairomones from tea aphids enhanced the responses to the plant–host complex. There was a significant, logistic dose–response relationship between the number of natural enemies responding and the odor stimulus concentration. Volatile components from the plant–host complex, obtained by air entrainment, were identified by their mass spectra and retention times and confirmed by comparison with standard samples. These were (Z)-3-hexen-1-ol, benzaldehyde, (E)-2-hexenal, (Z)-3-hexenyl acetate, ocimene, linalool, geraniol, indole, and (E)-2-hexenoic acid. The main components in a hexane rinse from tea aphid cuticle were benzaldehyde, undecane, 2,5-hexanedione, 2,5-dihydrothiophene, linalool, 4-methyl-octane, and eicosane, whereas the main components from an ether rinse were (E)-2-hexenoic acid, heptadecane, pentadecane, eicosane, tetratetracontane, and nonadecane. Benzaldehyde elicited the strongest responses from natural enemies in the olfactometer and the largest electroantennogram (EAG) responses. While the amount of odor was small, Coccinella septempunctata was slightly more sensitive than Chrysopa sinica and Aphidius sp. An increase in doses of benzaldehyde, (E)-2-hexenal, and (Z)-3-hexenyl acetate caused the EAG responses of each natural enemy to decrease. When the doses of (Z)-3-hexen-1-ol, linalool, and geranoil increased, EAGs of Chrysopa sinica and Aphidius sp. increased, but EAGs of Coccinella septempunctata decreased. When the dose of indole increased, EAGs of Coccinella septempunctata decreased, but those of Aphidius sp. increased. This study demonstrates that tea shoot–aphid complexes emit volatile synomones, while the odors from tea aphids, aphid cuticle extracts, and tea aphid honeydew contain kairomones, to which the natural enemies show a logistic dose–response.     

Antifeedant activity of extracts from neem,Azadirachta indica,to strawberry aphid,Chaetosiphon fragaefolii

Leaf disk choice test bioassays demonstrated that formulated neem seed oil (NSO) was equally deterrent to first- and third-instar nymphs and adult strawberry aphids,Chaetosiphon fragaefolii (Cockerell). Concentrations of NSO resulting in 50% feeding deterrence were approximately 1.1% for this species. The rapid disruption of aphid feeding (<1 hr) was not related to the presence of the limonoid azadirachtin, and deterrence likely results from the combined activity of several compounds. Activity toC. fragaefolii disappeared within 12–24 hr following application to strawberry in the greenhouse. NSO was deterrent to only half of the six aphid species tested. The antifeedant properties of neem do not appear to contribute significantly to the control of aphids and the viruses they transmit.     

Effect of Two Wheat Cultivars Differing in Hydroxamic Acid Concentration on Detoxification Metabolism in the AphidSitobion avenae

Hydroxamic acids (Hx) are wheat secondary metabolites conferring resistance for cereals against aphids. The activity of five enzymatic systems were evaluated in the aphid Sitobion avenae reared on the high-Hx wheat cultivar Chagual and the low-Hx wheat cultivar Huayún for 10 generations. Enzyme solutions were prepared from aphid homogenates and assayed for mixed function oxidases (including cytochrome P-450 monooxygenases and NADPH cytochrome c reductase), glutathione S-transferases, esterases, and catalase. Specific activities per aphid individual of cytochrome P-450 monooxygenases, NADPH cytochrome c reductase, glutathione S-transferases, and esterases were significantly increased in wheat cultivars relative to oat (only marginal increase of esterases in Chagual). Aphids fed on cv. Huayún showed an overall higher induction of enzymatic systems than those fed on cv. Chagual. Comparison of these results with reported effects of Hx on detoxifying enzymes in other insects, including aphids, support the hypothesis that these enzymatic pathways play an important role in the detoxification of toxic host-plant secondary metabolites.     

Effects of DIMBOA on several enzymatic systems in Asian corn borer,Ostrinia furnacalis (Guenée)

By using spectrophotometric analysis and isoelectric focusing electrophoresis (IEF), we found inhibition of acetylcholinesterase (AChE), general esterase activity (Est), and lipase activity in larvae of the Asian corn borer (ACB),Ostrinia furnacalis (Guenée), fed on cabbage dipped in DIMBOA (400 ppm). Amylase was not influenced, while activity of cytochrome P-450 monooxygenase and glutathloneS-transferase was increased. The results indicate that DIMBOA influences activity levels of some nervous system and detoxication enzymes and inactivates some hydrolysis enzymes.     

Hydroxamic Acid Content and Toxicity of Rye at Selected Growth Stages

Rye (Secale cereale L.) is an important cover crop that provides many benefits to cropping systems including weed and pest suppression resulting from allelopathic substances. Hydroxamic acids have been identified as allelopathic compounds in rye. This research was conducted to improve the methodology for quantifying hydroxamic acids and to determine the relationship between hydroxamic acid content and phytotoxicity of extracts of rye root and shoot tissue harvested at selected growth stages. Detection limits for an LC/MS-MS method for analysis of hydroxamic acids from crude aqueous extracts were better than have been reported previously. (2R)-2-β-d-Glucopyranosyloxy-4-hydroxy-(2H)-1,4-benzoxazin-3(4H)-one (DIBOA-G), 2,4-dihydroxy-(2H)-1,4-benzoxazin-3(4H)-one (DIBOA), benzoxazolin-2(3H)-one (BOA), and the methoxy-substituted form of these compounds, (2R)-2-β-d-glucopyranosyloxy-4-hydroxy-7-methoxy-(2H)-1,4-benzoxazin-3(4H)-one (DIMBOA glucose), 2,4-hydroxy-7-methoxy-(2H)-1,4-benzoxazin-3(4H)-one (DIMBOA), and 6-methoxy-benzoxazolin-2(3H)-one (MBOA), were all detected in rye tissue. DIBOA and BOA were prevalent in shoot tissue, whereas the methoxy-substituted compounds, DIMBOA glucose and MBOA, were prevalent in root tissue. Total hydroxamic acid concentration in rye tissue generally declined with age. Aqueous crude extracts of rye shoot tissue were more toxic than extracts of root tissue to lettuce (Lactuca sativa L.) and tomato (Lycopersicon esculentum Mill.) root length. Extracts of rye seedlings (Feekes growth stage 2) were most phytotoxic, but there was no pattern to the phytotoxicity of extracts of rye sampled at growth stages 4 to 10.5.4, and no correlation of hydroxamic acid content and phytotoxicity (I₅₀ values). Analysis of dose–response model slope coefficients indicated a lack of parallelism among models for rye extracts from different growth stages, suggesting that phytotoxicity may be attributed to compounds with different modes of action at different stages. Hydroxamic acids may account for the phytoxicity of extracts derived from rye at early growth stages, but other compounds are probably responsible in later growth stages.     

A Comparison of Semiochemically Mediated Interactions Involving Specialist and Generalist <Emphasis Type="Italic">Brassica</Emphasis>-feeding Aphids and the Braconid Parasitoid <Emphasis Type="Italic">Diaeretiella rapae</Emphasis>

Diaeretiella rapae, a parasitoid that predominately specializes in the parasitism of Brassica-feeding aphids, attacks Lipaphis erysimi, a specialist feeding aphid of the Brassicaceae and other families in the Capparales, at a greater rate than the generalist-feeding aphid, Myzus persicae. In this study, we investigated the orientation behavior of D. rapae to the volatile chemicals produced when these two aphid species feed on turnip (Brassica rapa var rapifera). We showed no significant preference orientation behavior to either aphid/turnip complex over the other. Isothiocyanates are among the compounds emitted by plants of the Brassicaceae in response to insect feeding damage, including by aphids. We assessed parasitoid orientation behavior in response to laboratory-formulated isothiocyanates. We tested two formulations and discovered significant orientation toward 3-butenyl isothiocyanate. We also assessed plant and aphid glucosinolate content, and showed large levels of glucosinolate concentration in L. erysimi, whereas there was little change in plant content in response to aphid feeding. Our results suggest that during the process of host location, similar cues may be utilized for locating L. erysimi and M. persicae, whereas the acceptance of hosts and their suitability may involve aspects of nonvolatile aphid chemistry.     

Change in acceptability of barley plants to aphids after exposure to allelochemicals from couch-grass (Elytrigia repens)

The response of the bird cherry-oat aphid, Rhopalosiphum padi, to barley plants was investigated following exposure of the plants to root allelochemicals from the aggressive weed couch-grass, Elytrigia (Agropyron) repens. Plants were treated either with root exudates from living couch-grass plants or with previously identified couch-grass root compounds [5-hydroxyindole-3-acetic acid, DL-5-hydroxytryptophan, L-5-hydroxytryptophan hydrate, and 6-hydroxy-1,2,3,4-tetrahydro--carboline-3-carboxylic acid (carboline)] either separately or in mixtures. In choice and no-choice settling tests, aphid acceptance of barley plants was significantly reduced following treatment with root exudates, and the carboline when tested alone or in combination with the other compounds. In contrast, the other compounds without the carboline were less active in reducing aphid acceptance. In a probing bioassay, individual substances were either neutral or stimulatory to aphids, indicating that the reduced settling was probably not due to direct effects on aphids, but rather due to effects on the plant. This was confirmed in olfactometer assays, in which aphids were repelled by odors from barley plants following treatment with a mixture containing all four chemicals.     

Relative importance of semiochemicals from first and second trophic levels in host foraging behavior ofAphidius ervi

The responses of femaleAphidius ervi to odors from a host food plant (Vicia faba), host aphids (Acyrthosiphon pisum), nonhost aphids (Aphis fabae), and aphid-plant complexes were investigated in a specially designed wind tunnel and a Y-tube olfactometer. In single-target (no-choice) and two-target (dual-choice) experiments, plant volatiles played a crucial role in the host foraging behavior ofA. ervi. The odor from theA. pisum-plant complex elicited the strongest responses byA. ervi females, followed by the odor from plants previously damaged by the feeding ofA. pisum. There was a significantly weaker response to odor fromA. pisum in the absence of the plant and to undamaged plants. Similarly, mechanically damaged plants and plants infested with the nonhost aphidA. fabae did not elicit strong responses. A plant that had been damaged byA. pisum and subsequently washed with distilled water was as attractive as an unwashed, previously infested plant.Aphidius ervi probably overcomes the reliability-detectability problem by selectively responding to herbivore-induced, volatile, semiochemical cues emitted by the first trophic level and by distinguishing between the volatiles induced by host and nonhost aphids.     

Responses of the Aphids Phorodon humuli and Rhopalosiphum padi to Sex Pheromone Stereochemistry in the Field

Gynoparous female and male damson-hop aphids, Phorodon humuli (Schrank), were caught in the field by water traps that were releasing the sex pheromone of this species, (1RS,4aR,7S,7aS)-nepetalactol. No behavioral activity was elicited by (4aS,7S,7aR)-nepetalactone, the major sex pheromone component of other aphid species such as Megoura viciae Buckton, even though olfactory cells were found in the secondary rhinaria on the third antennal segment of P. humuli that responded strongly to this compound. Gynoparous female P. humuli in the field responded less strongly to (1R,4aS,7S,7aR)-nepetalactol, the sex pheromone of the bird cherry-oat aphid, Rhopalosiphum padi (L.), than they did to the (4aR,7S,7aS)-nepetalactols, but males responded only to the latter. The (4aR,7S,7aS)-nepetalactone showed no electrophysiological activity so was not used in field trials. Releasing either the (4aS,7S,7aR)-nepetalactone or the (1R,4aS,7S,7aR)-nepetalactol with the (4aR,7S,7aS)-nepetalactols did not inhibit the response of P. humuli gynoparous females and males to the latter. Males of R. padi responded as strongly to the (4aR,7S,7aS)-nepetalactols as they did to (1R,4aS,7S,7aR)-nepetalactol. Males of P. humuli and R. padi responded positively to an increased concentration of the (4aR,7S,7aS)-nepetalactols released from two vials compared with that from a single vial, as did P. humuli (in one of two experiments) and R. padi to the (1RS,4aR,7S,7aS)- and (1R,4aS,7S,7aR)-nepetalactols when released together.     

Consumption and utilization of experimentally altered corn by southern armyworm: Iron,nitrogen, and cyclic hydroxamates

The effects of differential leaf water, leaf nitrogen and cyclic hydroxamate (DIMBOA) concentrations in corn seedlings were analyzed for a polyphagous insect, the southern armyworm (Spodoptera eridania Cram.). Six different combinations of nutrients and allelochemicals [DIMBOA = 2,4-dihydroxy-7-methoxy(2H)-benzoxazin-3(4H)-one] were generated using two corn genotypes (WF9 and CI3IA) and three fertility regimes (complete nutrient, Fe-deficient, and N-deficient solutions) in the University Biotron. Poorest larval growth was observed in the low-nitrogen treatments (1.2% and 1.7% leaf N) and was the result of both low consumption rates and high metabolic costs (low efficiency of conversion of digested food, ECD). Fastest growth rates were observed forthe larvae fed leaves from the high-nitrogen treatments (4.6% and 4.4% leaf N). It is noteworthy that these treatments also contained the highest concentration of cyclic hydroxamates, which are generally believed to be the primary defensive chemicals mediating resistance against the European corn borer,Ostrinia nubilalis (Hubner). If these hydroxamates do have any deleterious or costly effects (perhaps accounting for a large portion of metabolic expenditures), the high digestibility of the leaf tissue and the increased consumption rates more than compensate, resulting in rapid growth (growth rate = consumption rate × approximate digestibility × efficiency of conversion of the digested food). These studies illustrate that variation in key nutrients and allelochemicals within a single plant species (Zea mays L.) may have significantly different effects upon various potential leaf-chewing caterpillars, such as these armyworms versus corn borers (which cannot handle the cyclic hydroxamates, even if provided with young nutritious leaf tissues).     

Toxicokinetics of 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA) in the European corn borer,Ostrinia nubilalis (Hübner)

2,4-Dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA), the major hydroxamic acid present in corn, and its tritiated derivative, were prepared synthetically for use in the determination of the toxicokinetics of this insect deterrent in the European corn borer (ECB),Ostrinia nubilalis. In growth studies with DIMBOA (0, 0.05, 0.2, and 0.5 mg/g diet), the mean time to pupation and adult emergence were significantly lengthened by an increase in concentration. Pupal and adult weights, for both female and male, decreased with an increase in concentration. Increased larval and pupal mortality occurred at the highest concentration of DIMBOA. DIMBOA, at concentrations of 0.2 and 0.5 mg/g diet, resulted in a decrease in the number of egg masses produced per female, and at 0.5 mg/g diet, in a decrease in the number of eggs per egg mass. Larvae fed from the neonate stage on a diet containing 0.2 mg [³H]- + [¹H]DIMBOA/g diet showed an increase in the content of label from fourth to fifth instar, but levels declined at pupation and emergence. A large amount of the labeled compounds was excreted by the insect in the pupal case. In dose-related studies, both uptake and excretion increased with an increase in concentration of DIMBOA (0.05, 0.2, 0.4 mg/g diet), while a body burden (concentration in the tissues/concentration in the frass) of approximately 0.25 was maintained for all concentrations. At the highest dose of DIMBOA (0.4 mg/g), the ECB increased consumption, possibly to compensate for the toxic effects of the compound. In topical application studies, elimination of the labeled compound in the frass was rapid, reaching 65% by 4 hr and 88% by 48 hr. Accumulation of label in tissues other than hemolymph was low. The results show that the ECB does possess adaptive mechanisms to deal with the effects of this host-derived compound.     

Accumulation of Glucosinolates by the Cabbage Aphid Brevicoryne brassicae as a Defense Against Two Coccinellid Species

Brassica nigra plants, characterized by high levels of sinigrin, and artificial aphid diets to which sinigrin was selectively added were used to rear the crucifer specialist, Brevicoryne brassicae. Aphids were provided as a food source to two species of polyphagous ladybird, Adalia bipunctata and Coccinella septempunctata. First instar A. bipunctata were unable to survive when fed with B. brassicae reared on B. nigra or diets containing 0.2% sinigrin, but when fed with aphids reared on diets containing 0% sinigrin, survival rates were high. By contrast, first instar C. septempunctata were able to survive when fed with aphids reared on B. nigra or artificial diets containing up to 1% sinigrin. However, the presence of sinigrin in the aphid diet decreased larval growth and increased the time necessary for larvae to reach second instar for this species of ladybird. These results indicate that the presence of sinigrin in the diet of B. brassicae makes this aphid unsuitable as a food source for A. bipunctata but not for C. septempunctata, although for this ladybird species, there appear to be costs associated with feeding on aphids that contain this secondary metabolite.     

Methyl Salicylate,a Soybean Aphid-Induced Plant Volatile Attractive to the Predator <Emphasis Type="Italic">Coccinella septempunctata</Emphasis>

Induced volatiles provide a signal to foraging predatory insects about the location of their prey. In Iowa, early in the growing season of soybean, Glycine max, many predacious seven-spotted lady beetles, Coccinella septempunctata, were observed on plants with heavy infestations of soybean aphid, Aphis glycines. We studied whether the attraction of this beetle is caused by the release of specific volatile compounds of soybean plants infested by aphids. Volatile compounds emitted by soybean plants infested by aphids were compared with those of undamaged, uninfested, and artificially damaged plants. Gas chromatography–mass spectrometry analyses revealed consistent differences in the profiles of volatile compounds between aphid-infested soybean plants and undamaged ones. Significantly more methyl salicylate was released from infested plants at both the V1 and V2 plant growth stages. However, release patterns of two other induced plant volatiles, (d)-limonene and (E,E)-α-farnesene, differed between the two plant growth stages. Gas chromatographic–electroantennographic detection of volatile extracts from infested soybean plants showed that methyl salicylate elicited significant electrophysiological responses in C. septempunctata. In field tests, traps baited with methyl salicylate were highly attractive to adult C. septempunctata, whereas 2-phenylethanol was most attractive to the lacewing Chrysoperla carnea and syrphid flies. Another common lady beetle, the multicolored Asian lady beetle, Harmonia axyridis, showed no preference for the compounds. These results indicate that C. septempunctata may use methyl salicylate as the olfactory cue for prey location. We also tested the attractiveness of some selected soybean volatiles to alate soybean aphids in the field, and results showed that traps baited with benzaldehyde caught significantly higher numbers of aphids.     

Sex Pheromone of the Peach Aphid, Tuberocephalus momonis, and Optimal Blends for Trapping Males and Females in the Field

Chemical analysis of the volatiles released by sexual females (oviparae) of the peach aphid, Tuberocephalus momonis, identified two ubiquitous aphid sex pheromone components, (4aS,7S,7aR)-nepetalactone and (4aS,7S,7aR)-nepetalactol, in a ratio of 4 : 1. In field trials in Korea, employing traps releasing the two compounds in differing ratios, the most effective sex pheromone blend for trapping male T. momonis was found to be 85 : 15 nepetalactone–nepetalactol. Surprisingly, large numbers of presexual females (gynoparae) of this species were also collected when the catching rates were highest. In addition to T. momonis, over 20 other species of aphids were caught, particularly Myzus lythri, M. dycei, Lachnus tropicalis and M. persicae, in descending order of abundance.     

Distribution and Exudation of Allelochemicals in Wheat Triticum aestivum

Wheat allelopathy has potential for weed suppression. Allelochemicals were identified in wheat seedlings, and they were exuded from seedlings into agar growth medium. p-Hydroxybenzoic, trans-p-coumaric, cis-p-coumaric, syringic, vanillic, trans-ferulic, and cis-ferulic acids and 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA) were identified in both the shoots and roots of 17-day-old wheat seedlings and their associated agar growth medium. Wheat accessions with previously identified allelopathic activity tended to contain higher levels of allelochemicals than poorly allelopathic ones. The allelopathic compounds present in the shoots generally also were identified in the roots and in the agar medium. Allelochemicals were distributed differentially in wheat, with roots normally containing higher levels of allelochemicals than the shoots. When the eight allelochemicals were grouped into benzoic acid and cinnamic acid derivatives, DIMBOA, total coumaric, and total ferulic acids, the amount of each group of allelochemicals was correlated between the roots and the shoots. Most of the allelochemicals identified in the shoots and roots could be exuded by the living roots of wheat seedling into the agar growth medium. However, the amounts of allelochemicals in the agar growth medium were not proportional to those in the roots. Results suggest that wheat plants may retain allelochemicals once synthesized. The presence of allelochemicals in the agar growth medium demonstrated that wheat seedlings were able to synthesize and to exude phytotoxic compounds through their root system that could inhibit the root growth of annual ryegrass.