Evidence for volatile male-produced pheromone in banana weevilCosmopolites sordidus

Females of the banana weevil,Cosmopolites sordidus, were attracted to and made longer visits to live conspecific males, trapped volatiles from males, and dissected male hindguts in a still-air olfactometer. Male weevils were attracted to volatiles trapped from males and made longer visits to live males and volatiles from males. Live females, collected volatiles from females and female hindguts, elicited small or no behavioral responses from either sex. Electroantennogram (EAG) responses from both male and female antennae were elicited by collected volatiles from males and by dichloromethane extracts of male hindguts and bodies but not by surface washes of males. No significant EAG responses were given to equivalent material from females. It is therefore suggested that male banana weevils release an aggregation pheromone via their hindgut.     

Behavioral and electrophysiological responses of Aphids to host and nonhost plant volatiles

Alate and apterous virginoparae ofAphis fabae Scop, and alate virginoparae ofBrevicoryne brassicae (L.), walking in a linear track olfactometer, were attracted by odor from leaves of their host plants.A. fabae responded to odor from undamaged but not damaged bean leaves. Gynoparae (autumn migrants) ofA. fabae, however, did not respond to their host plant (spindle,Euonymus europaeus) odor. Odors of certain nonhost plants masked the attractiveness of the host plant leaves, but tansy (Tanacetum vulgare) and summer savory (Satureja hortensis) volatiles repelledB. brassicae andA. fabae, respectively. 3-Butenyl isothiocyanate attractedB. brassicae andLipaphis erysimi (Kalt.), the latter species being more sensitive in both behavioral and electrophysiological studies. Isothiocyanate receptors were found on the antennae ofA. fabae, which was repelled by these compounds, 4-pentenyl isothiocyanate being the most active.     

Behavioral responses of the parsnip webworm to host plant volatiles

Sixth instars of the parsnip webworm, Depressaria pastinacella, orient by olfaction to bud, male flowers, and female flowers of their primary host plant, Pastinaca sativa. Because octyl acetate and octyl butyrate are characteristic of tissues consumed by the sixth instar, we investigated the influence of these esters on webworm feeding behavior and chemo-orientation. Although octyl acetate and octyl butyrate are feeding deterrents, and octyl butyrate is an olfactory repellent, octyl acetate serves as an olfactory attractant. In olfactometers, webworms do not show a preference when given a choice between octyl acetate and host plant tissues. These findings suggest that octyl acetate is a sufficient cue for olfactory orientation. Such behavior may explain differences in the relative abundance of these esters observed among populations of wild parsnip under differential selection pressure from these insects.     

Behavioral and electrophysiological response of cabbage seed weevils (Ceutorhynchus assimilis) to conspecific odor

The responses ofCeutorhynchus assimilis Payk. (Coleoptera, Curculionidae) to the odor of overwintered and new generation weevils were studied using an olfactometer, choice tests in a laboratory cage, field tests using sticky traps baited with live weevils, and by electroantennograms (EAG's). Unmated male weevils and, to a lesser extent, female weevils of the overwintered generation were attracted to the odor of live unmated over-wintered female weevils. New generation weevils exhibited no behavioral response to conspecific odor. Male and female weevils of the overwintered generation exhibited positive EAGs to hexane extracts of overwintered female weevils, whereas EAGs of new-generation weevils of either sex were unresponsive to these extracts. This suggests that the unmated female weevils from the overwintered generation produce a volatile chemical or chemicals that attracts unmated male and female weevils. The new generation of female weevils does not produce this attractive chemical before overwintering, and male and female weevils of this generation can not detect the chemical(s) via their antennal chemoreceptors until they have undergone their overwintering period.     

Nitrogen and potassium fertilizer vs. nematode and weevil effects on yield and foliar nutrient status of banana in Uganda

Banana (Musa AAA) production is declining in central Uganda. A decline in soil fertility is often cited as one of the causes of the declining production. From 1996 to 1998, we studied banana yield responses to N and K fertilization at three sites in Uganda, where plantations ranged in age from 8 to 50 years. Phosphorus was applied to all treatments at 25 kg ha⁻¹, while N and K were applied (kg ha⁻¹) at 0N-0K, 0N-100K, 100N-0K, 100N-100K, and 100N-200K. Fresh fruit yields ranged from 4 to 67 Mg ha⁻¹ yr⁻¹. At one site the yield response to 100N-100K fertilization was significant (p < 0.01). The other sites showed positive but non-significant yield responses to N or NK fertilization. Economic analysis indicated that fertilizer use would be profitable only at the single site with significant yield increases. This site had negligible banana weevil (Cosmopolites sordidus (Germar)) and nematode (Radopholus similis; Helicotylenchus multicinctus) populations whereas the other sites were characterized by either severe nematode or weevil pressure. Fertilizer did not affect the damage caused by either nematodes or weevils. The best predictor of yield (p< 0.001, r ² = 0.75) was the number of functional leaves during anthesis. Measured soil nutrient parameters were highly variable but fell within sufficiency ranges. Foliar N contents were near diagnostic norms, but K levels fell below them, even when yields were satisfactory, suggesting that norms developed from other regions may not be suitable for Uganda. We conclude that in established plantations soil nutrient status is but one, and often a secondary, factor in the decline of banana yields in the central region of Uganda. Pest and disease infestations that result in reduced plantation productivity will need to be addressed before fertilizer use is likely to make a significant improvement in banana fruit yields. This revised version was published online in June 2006 with corrections to the Cover Date.     

Electroantennographic and behavioral responses of the sphinx moth Manduca sexta to host plant headspace volatiles

Coupled gas chromatography with electroantennographic detection (GC-EAD) using antennae of adult female Manduca sexta was employed to screen for olfactory stimulants present in headspace collections from four species of larval host plants belonging to two families: Solanaceae—Lycopersicon esculentum (tomato), Capiscum annuum (bell pepper), and Datura wrightii; and Martyniaceae—Proboscidea parviflora. Headspace volatiles were collected from undamaged foliage of potted, living plants. GC–EAD revealed 23 EAD-active compounds, of which 15 were identified by GC-mass spectrometry. Identified compounds included aliphatic, aromatic, and terpenoid compounds bearing a range of functional groups. Nine EAD-active compounds were common to all four host plant species: (Z)-3-hexenyl acetate, nonanal, decanal, phenylacetaldehyde, methyl salicylate, benzyl alcohol, geranyl acetone, (E)-nerolidol, and one unidentified compound. Behavioral responses of female moths to an eight-component synthetic blend of selected tomato headspace volatiles were tested in a laboratory wind tunnel. Females were attracted to the blend. A comparison of responses from antennae of males and females to bell pepper headspace volatiles revealed that males responded to the same suite of volatiles as females, except for (Z)-3-hexenyl benzoate. EAD responses of males also were lower for (Z)- and (E)-nerolidol and one unidentified compound. Electroantennogram EAG dose–response curves for the 15 identified EAD-active volatiles were recorded. At the higher test doses (10–100 g), female antennae yielded larger EAG responses to terpenoids and to aliphatic and aromatic esters. Male antennae did respond to the higher doses of (Z)-3-hexenyl benzoate, indicating that they can detect this compound. On the basis of ubiquity of the EAD-active volatiles identified to date in host plant headspace collections, we suggest that M. sexta uses a suite of volatiles to locate and identify appropriate host plants.     

Olfactory Responses of Banana Weevil Predators to Volatiles from Banana Pseudostem Tissue and Synthetic Pheromone

As a response to attack by herbivores, plants can emit a variety of volatile substances that attract natural enemies of these insect pests. Predators of the banana weevil, Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae) such as Dactylosternum abdominale (Coleoptera: Hydrophilidae) and Pheidole megacephala (Hymenoptera: Formicidae), are normally found in association with weevil-infested rotten pseudostems and harvested stumps. We investigated whether these predators are attracted to such environments in response to volatiles produced by the host plant, by the weevil, or by the weevil–plant complex. We evaluated predator responses towards volatiles from banana pseudostem tissue (synomones) and the synthetic banana weevil aggregation pheromone Cosmolure+ in a two-choice olfactometer. The beetle D. abdominale was attracted to fermenting banana pseudostem tissue and Cosmolure+, whereas the ant P. megacephala was attracted only to fermented pseudostem tissue. Both predators were attracted to banana pseudostem tissue that had been damaged by weevil larvae irrespective of weevil presence. Adding pheromone did not enhance predator response to volatiles from pseudostem tissue fed on by weevils. The numbers of both predators recovered with pseudostem traps in the field from banana mats with a pheromone trap were similar to those in pseudostem traps at different distance ranges from the pheromone. Our study shows that the generalist predators D. abdominale and P. megacephala use volatiles from fermented banana pseudostem tissue as the major chemical cue when searching for prey.     

Development of an Efficient Pheromone-Based Trapping Method for the Banana Root Borer <Emphasis Type="Italic">Cosmopolites sordidus</Emphasis>

The banana root borer Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae) is a major pest of bananas throughout the world. Chemical control is both undesirable and expensive, where biological control alternatives are limited, and pheromone-based trapping results in low captures. In this study, several important factors that affect pheromone-based catches, such as trap type, trap dimensions, and color and position of the traps, were optimized. Ground traps were found to be superior to ramp and pitfall traps, and larger traps (40 × 25 cm and above) were more efficient than smaller ones (30 × 15 cm). In a color-choice test, the banana weevil clearly preferred brown traps over yellow, red, gray, blue, black, white, and green, with mahogany being more attractive than other shades of brown. In addition, pheromone baited ground traps positioned in the shade of the canopy caught significantly more adults than those placed in sunlight. Therefore, mahogany-brown ground traps 40 × 25 cm appear to be the most efficient at catching C. sordidus adults and have the greatest potential for use in mass trapping and programs for eradication of this pest.     

Chemosensory responses of cowpea weevil,Callosobruchus chinensis to an aquatic weed,water hyacinth,Eichhornia crassipes (Mart) Solms

The behavioral responses of cowpea weevil,Callosobruchus chinensis to the leaf extracts of the aquatic weed, water hyacinth,Eichhornia crassipes (Mart) Solms. was evaluated by three different methods. The petroleum ether-soluble fraction ofE. crassipes induced high weevil attraction in all methods employed. The olfactometer assay provided a reading on weevil chemosensory responses, whereas the free-choice tests measured response and preference of weevils to the untreated or extract-treated food grain of their choice over a longer period. The extracts were also presented under no choice test conditions to assay them for the presence of oviposition stimulants. In all these methods the extracts of water hyacinth evoked a quick directional response. Another interesting feature is that the extract, besides producing attraction, also caused mortality of the insects.     

Orientation of boll weevil,Anthonomus grandis boh. (Coleoptera: Curculionidae), to pheromone and volatile host compound in the laboratory

Behavioral responses of male and female boll weevils to the aggregation pheromone, grandlure, and the major volatile of cotton, -bisabolol, were investigated using a new dual-choice olfactometer. Dosage-response experiments revealed both males and females to be attracted by the aggregation pheromone at the 1.0 g dosage. However, only males were attracted to -bisabolol (1.0 g). Both sexes were repelled by the highest dosage of-bisabolol tested (10 g). In preference experiment, males chose grandlure over -bisabolol, while both sexes chose the combination of grandlure + -bisabolol over -bisabolol alone. There was some evidence for synergism between pheromone and plant odor for the females. The results correlate well with previous electrophysiological and behavioral experiments.Mention of a trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products or vendors that may also be suitable.     

Antennal perception of oilseed rape,Brassica napus (Brassicaceae), volatiles by the cabbage seed weevilCeutorhynchus assimilis (Coleoptera,Curculionidae)

The response of theCeutorhynchus assimilis antenna to volatiles in air entrainment-derived extracts of oilseed rape,Brassica napus, was studied using coupled gas chromatography (GC)-electroantennography (EAG) and coupled GC-single cell recording (SCR). By means of these techniques and coupled gas chromatography-mass spectrometry (GC-MS), 25 active compounds were identified, including isoprenoids and compounds derived from fatty acids and amino acids. Some of the latter, the isothiocyanates and goitrin, and probably indole and benzyl cyanide, are catabolites of glucosinolates. The electrophysiological activity of the identified compounds was confirmed by EAG using a physiologically discriminating dose, and by SCR studies. The importance of the combined use of the EAG and SCR techniques was demonstrated, since specific olfactory cells were located for five compounds that did not elicit significant EAG responses. The majority of the olfactory cells from which single cell recordings were obtained showed very high specificity, and in numerous recordings there were consistent pairings of specific cell types.     

Antennal olfactory response of boll weevil to grandlure and vicinal dimethyl analogs

Electroantennogram techniques were used to elucidate antennal olfactory response of male and female boll weevils to a dilution series of grandlure, its components, and some vicinal dimethyl analogs. At higher concentrations, response to the mixture of the two aldehyde components of grandlure was significantly higher than to the two alcohol components. Only one vicinal dimethyl analog elicited a significantly higher response than the control. There were no significant differences in response due to sex over all compounds.Texas Agricultural Experiment Station paper number 16382. Research supported by McIntire-Stennis project 1525. All programs of the Texas Agricultural Experiment Station are available without regard to race, sex, color or religion.     

Attraction of adult sweet potato weevils,Cylas formicarius elegantulus (Summers), (Coleoptera: Curculionidae), to sweet potato leaf and root volatiles

A dual-choice olfactometer was developed to study the responses of sweet potato weevils,Cylas formicarius elegantulus (Summers), to volatiles from the sweet potato,Ipomoea batatas (L.) Lam. Both males and females were attracted by volatiles from sweet potato leaves and a methylene chloride leaf extract. Females, but not males, responded to volatiles from storage roots and a methylene chloride root extract. Leaves and storage roots from four sweet potato cultivars (Centennial, Jewel, Resisto, and Regal) were attractive to female weevils; however, the attractant response varied with cultivar. GC profiles from leaf and root extracts, and GC-MS analysis of leaf extract, for Jewel cultivar enabled the volatile peaks to be identified as sesquiterpenes.     

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.     

Olfactory responses of adultTribolium castaneum (Herbst), to volatiles of wheat and millet kernels,milled fractions,and extracts

Olfactory responses of 72- to 96-hr-old, 24-hr starved adult male, female, or mixed-sex groups of the red flour beetle,Tribolium castaneum (Herbst), to volatiles from wheat and millet kernels, certain milled fractions, and solvent extracts were recorded by using a lightsensitive apparatus. Wheat-germ volatiles from intact germ or solvent extracts were generally more attractive than volatiles of wheat endosperm or wheat bran. Volatiles of whole wheat kernels were the least attractive among test materials of wheat origin, whereas there were no significant differences among whole wheat flour, germ, and endosperm. Wheat-germ extracts, however, were more attractive than were extracts of the other fractions. Whole millet flour or fermented millet flour volatiles were more attractive than those from whole millet kernels or millet starch. Beetles reached maximum responsiveness to grain volatiles by 72–96 hr after adult ecdysis. Groups of virgin female beetles generally were more responsive than male or mixed-sex groups to volatiles of substances tested.Contribution 82–137-j, Department of Entomology, Kansas Agricultural Experiment Station (Project Hatch 322), Kansas State University, Manhattan, Kansas.     

Behavioral and neurosensory responses of the boll weevil,Anthonomus grandis Boh. (Coleoptera: Curculionidae), to fluorinated analogs of aldehyde components of its pheromone

Competitive field tests with -fluorinated analogs of compounds III and IV (III--F and IV--F, respectively) of the boll weevil,Anthonomus grandis Boh., aggregation pheromone showed these compounds, when combined with the other pheromone components [(±)-I and II], to be as attractive as grandlure [(+)-I, II, and III+IV]. Dose-response curves constructed from electroantennograms of male boll weevils to serial stimulus loads of III, IV, III--F, IV--F, and the corresponding acyl fluorinated analogs (III-acyl-F and IV-acyl-F) showed the -fiuorinated analogs to be as active as the pheromone components (threshold=0.1 g), while the acyl fluorinated analogs had a 10-100 x higher threshold (=1-10 g). Single-neuron recordings showed that IV neurons and II neurons (Dickens, 1990) responded to IV--F and III--F, respectively, while IV-acyl-F and III-acyl-F were inactive. Since a previous study showed compounds I, II, and IV to be essential for behavioral responses in the field, it seems likely that the activity of the -fluorinated analogs observed here is due to the stimulation of IV neurons by IV--F as indicated in single neuron recordings.     

Grain weevil,Sitophilus granarius (L.): Antennal and behavioral responses to male-produced volatiles

Coupled GC-EAG techniques have been applied to the study of volatiles from the grain weevil,Sitophilus granarius. for the first time. The size of EAG response was independent of the sex of the responding insect but was consistently larger to extracts of males than those of females. This difference was reflected in a behavioral preference for the male extracts by mated adults of both sexes tested together and virgin adults of both sexes tested separately. The GC-EAG results provide evidence for two materials that are released specifically by the males. Using circular dichroism. one has been found to be identical stereochemically with the (2S,3R)-sitophilate reported by others as the aggregation pheromone in a different strain. This enhances the prospects for the development of a single pheromone lure that would be generally applicable whatever the origin of the strain. The small amount of sitophilate found in the males suggests that it is not stored in large amounts. The other material, present in such a small amount that it has yet to be fully characterized, elicits a higher antennal activity than sitophilate and may have a significant role to play in enhancing the trap catch of this economically important pest.     

Electroantennogram responses of the cabbage seed weevil,Ceutorhynchus assimilis,to oilseed rape,Brassica napus ssp.Oleifera,volatiles

Electroantennograms (EAGs) were recorded from male and female cabbage seed weevils (Ceutorhynchus assimilis Payk.) in response to volatiles isolated and identified from the odor of oilseed rape (Brassica napus ssp.oleifera DC. cv. Ariana). Relatively large EAGs were obtained on stimulation with volatiles produced by the oilseed rape crop at the time when seed weevils were actively searching for host plants. Artificial rape odor without certain key volatile compounds was in most cases significantly less stimulatory than odor containing these volatiles. There were significant differences in the EAG response of the sexes ofC. assimilis to the green leaf volatiles of oilseed rape and several terpenes present in rape flower odor. The importance of the qualitative and quantitative composition of host-plant odor in host location byC. assimilis is discussed.     

Behavioral and electrophysiological responses of Arhopalus tristis to burnt pine and other stimuli

The exotic longhorn beetle Arhopalus tristisis a pest of pines, particularly those damaged by fire, and a major export quarantine issue in New Zealand. Actinograph recordings of caged individuals showed that males and females were most active from dusk to midnight. Olfactometer experiments indicated that females moved upwind toward odors from burnt pine (80%, N= 75), compared to unburnt pine (20%). Oviposition choice tests showed that eggs were predominantly laid on burnt logs (79%, N= 20), compared to unburnt logs. Beetles were trapped by funnel traps baited with burnt (mean catch per trap 7.8) and unburnt (mean catch 4.1 per trap) pine bark from inside a screen cage (4 × 3 m), while unbaited traps had a mean catch 0.1 beetles (N= 8 replicates). The treatment of burnt pine bark with a 1:1:2 mixture of green leaf volatiles (E)-2-hexen-1-ol and (E)-2-hexenal) in mineral oil as a repellent reduced trap catch by fivefold in a similar experiment (mean catches of 1.2 beetles per trap to burnt pine bark plus repellent treatment and 6.2 beetles per trap to burnt pine bark alone). The treatment of burnt pine bark with this solution also reduced oviposition by 98.5% (mean eggs per log of 11.1 on burnt pine and 0.3 on burnt pine plus repellent), indicating that oviposition cues have the potential to be significantly disrupted. The electrophysiological responses of adult beetles were recorded to a range of odorants. Normalized responses to monoterpenes known to occur in Pinus radiataranged from about 20 to about 150, with -terpineol giving the greatest responses in both sexes. Green leaf volatiles also gave high responses. The potential exists to improve the management of this insect using chemical cues in various ways.     

Electroantennogram responses ofTrirhabda bacharides (weber) (Coleoptera: Chrysomelidae) to plant volatiles

Electroantennograms (EAGs) were recorded fromTrirhabda bacharides Le Conte (Coleoptera: Chrysomelidae) females to 28 odorants and volatiles emanating from leaves of 26 plant species including three known host plants,Baccharis halimifolia L.,B. neglecta Britt, andB. salicina T. & G. (Compositae: Asterae). Antennal receptors were highly responsive to components of the green leaf volatile complex, especially 6-carbon saturated and monounsaturated alcohols andtrans-2-hexenal. EAGs elicited by heptanal were greater than those elicited by any other saturated aldehyde. Oxygenated monoterpenes were more active than monoterpene hydrocarbons. -Bisabolol was the most active sesquiterpene. In general, EAGs to volatiles emanating fromBaccharis and other composite species were greater than those elicited by species from other families. Furthermore, when one considers only plants occurring in the environs ofT bacharides hostB. neglecta, EAGs elicited by its host plant were at least two times greater than those elicited by other coinhabiting plants.Mention of a trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products or vendors that may also be suitable.