Flight-mediated attraction ofBiprorulus bibax breddin (Hemiptera: Pentatomidae) to natural and synthetic aggregation pheromone

The attraction of female spined citrus bugs,Biprorulus bibax, to natural and synthetic aggregation pheromone was studied using an olfactometer and a large flight cage. No locomotory response by postdiapause, prereproductive females to heptane extracts of male dorsal abdominal glands (DAGs) (site of pheromone production) was recorded in the olfactometer study. However, postdiapause, prereproductive females showed significant attraction to sites baited with DAG extracts in the flight cage (1.9–3.0 times that of unbaited sites). Prereproductive and reproductive females showed greatest attraction to sites baited with a synthetic blend of pheromone [(3R,4S,1E-3,4-bis(1-butenyl)tetrahydro-2-furanol, linalool, farnesol, nerolidol] (2.3–4.7 times the attraction of unbaited sites). Females also responded significantly to sites baited with the hemiacetal major component alone (1.7–2.2×). Diapausing females collected from fall populations did not respond to natural or synthetic pheromone baits. Potential applications of the synthetic aggregation pheromone are discussed with respect toB. bibax management.     

Male-produced aggregation pheromone of the maize weevil,Sitophilus zeamais,and interspecific attraction between threeSitophilus species

A dual-choice pitfall bioassay was used to demonstrate the existence of a male-produced aggregation pheromone in the maize weevil. Both males and females showed a highly significant preference for extracts of disks exposed to wheat-feeding males over extracts of disks exposed to wheat kernels only. Neither sex responded significantly to extracts from females. Mating did not reduce pheromone release by males. Response by virgin females to pheromone was significantly higher than response by mated females, but males of either mating status responded equally well. There was no apparent daily periodicity in the responsiveness to pheromone. Rice and maize weevils showed a strong interspecific cross-attraction. Granary weevils of both sexes responded well to maize weevil extracts, but only females showed a significant response to rice weevil extracts. Neither maize nor rice weevils responded significantly to granary weevil extracts.     

Inhibition of the attractant pheromone response inIps pini andI. paraconfusus (Coleoptera: Scolytidae): Field evaluation of ipsenol and linalool

Linalool, a compound from maleI. pini, previously suggested as an inhibitor forI. paraconfusus, has no obvious effect on the response ofI. paraconfusus to ponderosa pine bolts containing maleI. paraconfusus. I. pini from California and New York equally inhibit the response ofI. paraconfusus to maleI. paraconfusus. Ipsenol, one component of the attractant pheromone ofIps paraconfusus, inhibits attacks byIps pini on ponderosa pine logs baited with maleI. pini. The concentration of ipsenol used appears to be critical for effective suppression of attacks.This investigation was supported in part by a grant from the U.S. Forest Service.     

Response ofDendroctonus brevicomis andIps paraconfusus (Coleoptera: Scolytidae) to combinations of synthetic pheromone attractants and inhibitors verbenone and ipsdienol

DendroctonusBrevicomis andIps paraconfusus are sympatric bark beetle species colonizingPinus ponderosa in western North America. Interspecific and intraspecific competition for resources is, in part, mediated through semiochemicals. The response ofD. brevicomis to its attractant pheromone was significantly reduced by simultaneous release of either verbenone or racemic ipsdienol. Trap catch was significantly further reduced by including both inhibitors with the attractant pheromones. However, although the response ofIps paraconfusus was significantly inhibited with the addition of either verbenone or racemic ipsdienol, both compounds together did not result in a significant further reduction in trap catch. There was a trend for greater reduction in response ofD. brevicomis to attractant pheromones with increased release rates of either 69% (+)-/31% (–)-verbenone or 84% (–)-/16% (+)-verbenone. Response of associates to attractants and inhibitor combinations was also determined.     

Calculated effect of pulsed pheromone release on range of attraction

In order to assess the possible function of pulsed release of windborne pheromones, calculations were performed to predict the concentration pattern downwind of the source. Two patterns of pulsed releases (sinusoidal and instantaneous) were compared to a constant release pattern. In all three cases, the average rate of release was the same. Assuming the receiving animal needs only a momentary exposure to concentrations above threshold to respond, it is clear that sinusoidal release has a greater distance of detection than constant release and that instantaneous release is even better. The relative magnitude of the increase in range of detection depends on the ratio of average release rate to threshold concentration. Pulsed releases have a greater advantage when the threshold is high and the range of attraction is inherently short. Under these conditions, sinusoidal release can double the range of attraction and instantaneous release can increase it 10-fold. In contrast, with a low threshold and consequent long range of attraction, the pulsed patterns are lost and the increase in range is insignificant. Several testable predictions are derived from the hypothesis that the primary function of pulsed release is to extend the range and time over which a given quantity of pheromone can act.     

Field attraction ofBiprorulus bibax Breddin (Hemiptera: Pentatomidae) to synthetic aggregation pheromone and (E)-2-hexenal,a pentatomid defense chemical

A synthetic blend of the aggregation pheromone [(3R,4S,1E)-3,4-bis(1-butenyl)tetrahydro-2-furanol, linalool, farnesol, and nerolidol] of the spined citrus bug,Biprorulus bibax, and the pentatomid defense chemical, (E)-2-hexenal, both attracted adultB. bibax to individual trees in citrus orchards. Lemon trees containing single glass vials with aggregation pheromone or (E)-2-hexenal were colonized by significantly greater numbers of reproductiveB. bibax than unbaited trees. There was no significant difference between the treatments and bug recruitment was not improved by using both treatments.B. bibax did not enter cylinder/funnel traps baited with aggregation pheromone but colonized trees containing the traps. Orange or lemon trees containing aggregation pheromone on orchard perimeters recruited significantly larger populations of emigrating, nonreproductiveB. bibax during fall than untreated trees. Nonreproductive bugs were not attracted to trees containing (E)-2-hexenal. The potential for using these semiochemicals as management tools forB. bibax is discussed.     

Male olive fruit fly attraction to synthetic sex pheromone components in laboratory and field tests

Male olive fruit fly attraction to the four synthetic components of the female sex attractant pheromone was studied under laboratory and field conditions. In laboratory tests males responded to all four components tested separately. Component I, (1,7-dioxaspiro[5.5]undecane) was more attractive than any of the remaining three components alone, but a combination of all four was more attractive than component I alone. In field tests with polyethylene vials as pheromone dispensers, the complete mixture, although not statistically significant, was constantly more attractive to males than component I alone. A tendency of enhancement of attraction of component I by combining it with component II (-pinene) or III (n-nonanal) was also observed. In field tests with rubber septa as pheromone dispensers only component I was attractive. Mixtures containing component I were also attractive but not more attractive than component I alone. Evaporation rate and ratio of components as they come out of the dispenser appear to be critical for male response.     

Interactions between components of the aggregation pheromone during chemotaxis by the bark beetleIps paraconfusus

The male-produced aggregation pheromone inIps paraconfusus is composed of three compounds. Female bark beetles were exposed to combinations of these compounds, presented as point sources in an enclosed, circular arena. By itself,cis-verbenol (cV) had no effect on the number of beetles that reached the source. Either ipsenol (Ip) alone or ipsdienol (Id) alone strongly increased the number that reached the source, with Id producing a dose-response curve with a much steeper slope. cV moved the onset of the response to Id to higher doses of Id, but the response rose more rapidly after onset than when cV was absent. Overall, cV inhibited the effect of Id except at the highest dose. cV affected the onset of the response to Ip little or none, but strongly increased the slope of the response, synergizing the effect of Ip. The responses to combinations of Id and Ip were related to the log of a linear combination of their doses. The results are consistent with a model where Id and Ip act at a single site of action, but with different potencies, while cV appears to modify the effects of Id and Ip, rather than affecting the site of action directly.     

Field response of spruce bark beetle,Ips typographus,to aggregation pheromone candidates

Six compounds previously identified from hindguts of unmated maleIps typographus (L.) during host colonization: 2-methyl-3-buten-2-ol (MB),cis-verbenol (cV),trans-verbenol (tV), myrtenol (Mt),trans-myrtanol (tM), and 2-phenylethanol (PE), were tested for their attractivity in the field with a subtractive method. The amounts of MB and cV released from a pipe trap were similar to those given off from the commercial bait Ipslure as well as that from a Norway spruce tree,Picea abies (L.) Karst., under mass attack. The blend of the compounds became nonattractive when either MB or cV was subtracted, while subtraction of any of the other four compounds had no effect. Addition of ipsdienol (Id) to the blend did not significantly increase the attraction. In a second comparative test, the addition of three compounds as a group (tV + Mt + PE) to MB + cV again had no effect on the attraction, but the addition of Id increased the catch somewhat. Addition of host logs to a bait releasing MB + cV at a rate lower than in previous experiments did not influence the attraction to pipe traps. Sticky traps containing natural pheromone sources (50 males in a log), which released 1–5 mg/day of MB as determined by aerations with deuterated MB as internal standard, were less attractive than a synthetic source releasing similar amounts of MB.Coleoptera: Scolytidae.This study was made within the Swedish project Odour Signals for Control of Pest Insects.     

Sex-specific responses to aggregation pheromone Regulation of colonization density in the bark beetleIps paraconfusus

About equal numbers of each sex of flyingIps paraconfusus Lanier (Coleoptera: Scolytidae) were caught on traps several meters downwind from a male-infested ponderosa pine log releasing pheromone while a significantly different ratio of over four times more females than males were caught at the pheromone source. Females oriented directly to higher concentrations of colonizing males in a felled tree while males tended to land on the host in adjacent uncolonized areas. The attraction response of walking males to a 1∶1∶1 mixture of the synthetic pheromone components ispenol-ipsdienol-cis-verbenol was reduced progressively at higher concentrations while female response continued to increase. These responses may function to regulate density of colonization and limit intraspecific competition.     

Response ofDendroctonus brevicomis Le Conte (Coleoptera: Scolytidae) to different release rates and ratios of aggregation semiochemicals and the inhibitors verbenone and ipsdienol

The response of western pine beetle,Dendroctonus brevicomis Le Conte, to different release rates of the aggregation semiochemicals (attractants)exo-brevicomin, frontalin, and myrcene and the inhibitors verbenone and ipsdienol was investigated. Release rates of verbenone ranging from 0.18 mg/ 24 hr to 1.2 mg/24 hr did not result in significant reductions in mean trap catch ofD. brevicomis. In contrast, very low release rates of ipsdienol (0.02–0.4 mg/24 hr) significantly reduced trap catch compared to controls. The combination of verbenone and ipsdienol, released at rates above 0.09 and 0.02 mg/24 hr, respectively, resulted in significantly lower trap catches ofD. brevicomis in attractant-baited traps. Results of an experiment testing a factorial combination of different release rates of verbenone and attractants suggest that response is not ratio-specific. The response curve ofD. brevicomis to the levels of verbenone was similar across all levels of attractants, while the response to equivalent ratios of attractants to verbenone was not similar, suggesting that the behavior of the beetles is primarily influenced by the absolute release rate of verbenone.     

Inhibition of beta-amyloid aggregation and neurotoxicity by complementary (antisense) peptides

Complementary peptides are coded for by the nucleotide sequence (read 5'-->3') of the complementary strand of DNA. By reading the sequence of complementary DNA in the 3'-->5' direction, alternative complementary peptides may be derived. We describe the derivation, testing and analysis of six complementary peptides designed against beta-amyloid peptide 1-40 (Abeta, 40). Data is presented to show that one peptide, designated 3' -->5' betaCP1-15, binds specifically to Abeta 1-40, and inhibits both fibrilisation and neurotoxicity in vitro. This suggests that complementary peptides could be useful leads for drug discovery, especially where diseases of protein misfolding are concerned.     

Photoisomerization of antiaggregation pheromone verbenone: Biological and practical implications with respect to the mountain pine beetle,Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

Release of the antiaggregation pheromone, verbenone, at 3.8 mg/ day from a concentrated source within a multiple-funnel trap completely inhibited response by the mountain pine beetle (MPB),Dendroctonus ponderosae Hopkins, to attractive semiochemical lures. When aerial applications were simulated and verbenone was released at the same rate from beads lying in a 2×2-m area on the forest floor 15–35 cm below a trap, the response of the MPB was inhibited by only 50%. This reduced inhibition may be explained in part by the photoisomerism of verbenone. When exposed to full sunlight on two occasions, the times required for 50% of verbenone vapors to be converted to chrysanthenone were 75 and 100 min, respectively. Trap and tree-baiting experiments indicated no biological activity of chrysanthenone. Rapid photoisomerization could reduce the concentration of verbenone below biologically active levels and would allow the MPB to colonize trees close to already occupied hosts, contributing to the characteristic clumped distribution of MPB attack. The rate of verbenone photoisomerization may vary according to geographic location, stand elevation and density, and should be considered before verbenone is applied to control the MPB and other bark beetles.     

Influence of aggregation inhibitors (verbenone and ipsidenol) on landing and attack behavior ofDendroctonus brevicomis (Coleoptera: Scolytidae)

The influence of the aggregation inhibitors verbenone and ipsdienol on the response of western pine beetle,Dendroctonus brevicomis, to attractive host trees was investigated. Paired ponderosa pine trees (Pinus ponderosa) were baited with aggregation semiochemicals to stimulate mass attack. One tree in each pair received an inhibitor treatment consisting of five sets of two verbenone and two ipsdienol dispensers spaced 1 m apart vertically along the tree bole. Beetle landing was monitored with sticky traps on the tree bole, and attack density was assessed from bark samples removed four or seven days after baiting. The inhibitor treatment resulted in a significant reduction of both the numbers of beetles landing on trees and the density of attacking beetles compared to control trees (without inhibitors). The ratios of beetle landing density to attacking density were not different between inhibitor-treated and control trees, nor were the vertical distributions of beetles landing or attacking, suggesting that beetle behavior was primarily influenced at a longer range, prior to landing on the tree. Although the application of verbenone and ipsdienol did not preventD. brevicomis from attacking baited trees, our results suggest that when applied to unattacked (and unbaited) trees, their effectiveness at reducing the attack pressure might allow trees having a certain amount of resistance to survive attack by pioneer beetles.     

Conversion of verbenols to verbenone by yeasts isolated fromDendroctonus ponderosae (Coleoptera: Scolytidae)

A variety of symbionts associated with bark beetles are capable of producing compounds that are used as pheromones by their hosts. We report that two yeasts associated withDendroctonus ponderosae Hopkins,Hansenula capsulata Wickerham, andPichiapinus (Hoist) Phaff, are capable of convertingcis- andtrans-verbenol efficiently into verbenone.trans-Verbenol, which is produced by femaleD. ponderosae, acts as an aggregation pheromone for this scolytid, while verbenone, which other studies have indicated that microbe-reducedD. ponderosae are incapable of producing, acts as an antiaggregation pheromone.D. ponderosae appears to rely primarily on microbial symbionts for terminating aggregation and mass attack on individual host trees.Research supported in part by the Natural Sciences and Engineering Research Council, Canada, Operating Grant A3881, and Strategic Grant G1039.     

Effect of foliage proximity on attraction ofChoristoneura occidentalis andC. retiniana (Lepidoptera: Tortricidae) to pheromone sources

Pheromone-baited traps located close to both host and nonhost crowns were more attractive than traps located between crowns for bothC. occidentalis Freeman andC. retiniana (Walsingham) at both 10 m and at 1.5 m above the ground. At 10 m height, traps located in host foliage were more attractive than those located in nonhost foliage, but at 1.5m height there was no significant difference. These results were obtained for both dense and sparse populations ofC. occidentalis and sparse populations ofC. retiniana. We conclude that the tree species on which a virgin female is located is not an important factor restricting mating between closely relatedChoristoneura spp. Also, the tree species on which a trap is located may not be an important factor that must be standardized in developing pheromone monitoring systems forC. occidentalis andC. retiniana.     

Identification and biosynthesis of an aggregation pheromone of the storage mite Chortoglyphus arcuatus

In an effort to identify new pheromones from mites, the headspace of undisturbed colonies of the storage mite Chortoglyphus arcuatus was analyzed by GC-MS by use of a closed-loop stripping apparatus (CLSA) or solid-phase microextraction (SPME). The major compound emitted from the mites is (4R,6R,8R)-4,6,8-trimethyldecan-2-one (4R,6R,8R-8). The structure was elucidated by analysis of the mass spectrum, synthesis of authentic samples, and gas chromatography on a chiral phase. Bioassays show that this compound, for which we propose the trivial name chortolure, is an aggregation pheromone for both sexes of this species. Several related compounds are released in smaller amounts by the mites. The alarm pheromones of these mites, neral and geranial, can only be found in total extracts of the mites, in which 8 occurs only in minute amounts. The method of sampling is therefore crucial for pheromone identification. Feeding experiments with deuterated propionate showed that chortolure is a polyketide, formed by successive addition of four propionate units to an acetate starter.     

Interruption of response ofDendroctonus brevicomis to its attractive pheromone by components of the pheromone

The catch of the western pine beetle,Dendroctonus brevicomis, at an attractive source of racemicexo-brevicomin, racemic frontalin, and myrcene was reduced by surrounding the source with a grid of 48 stations releasing all three compounds together, orexo-brevicomin alone or myrcene alone. Each compound was released at the rate of 2 mg/24 hr/station. The catch at an attractive bolt cut from a tree being colonized byD. brevicomis was not reduced byexo-brevicomin, but was reduced by the combination ofexo-brevicomin, frontalin, and myrcene in one of two tests. When a transect of traps was placed across a 0.81-hectare plot at six of the 48 stations releasing all three compounds, more beetles were caught at outer than at inner traps. More beetles were caught at unbaited traps on trees in a plot when the three compounds were released than when onlyexo-brevicomin or no compounds were released. A few trees were attacked byD. brevicomis in some of the plots. The antiattractant verbenone released from 48 stations at the rate of 4 mg/24 hr/station did not reduce the catch at an attractive tree bolt.Coleoptera: Scolytidae.These studies were supported by the Forest Service, U.S. Department of Agriculture, and in part by grants to the University of California from the U.S. National Science Foundation and U.S. Environmental Protection Agency (NSF GB-34718/BMS 75-04223) and the Rockefeller Foundation. The findings, opinions, and recommendations are not necessarily those of the funding agencies. Mention of commercial enterprises or products is only for information.     

Ester components of aggregation pheromone ofDrosophila virilis (Diptera: Drosophilidae)

The male-produced aggregation pheromone ofDrosophila virilis was found to contain five ester components, in addition to a previously identified hydrocarbon, (Z)-10-heneicosene (Z10–21). The five esters were: the methyl, ethyl, and 1-methylethyl (isopropyl) esters of 2-methyl-(E)-2-butenoic (tiglic) acid and the methyl and ethyl esters of hexanoic acid. The esters were not detected in females. Each ester was active by itself in laboratory bioassay tests, and each increased the number of flies responding toZ10–21 ca. 4–5 times. In comparisons among the five esters at 10 ng per compound, ethyl tiglate was the most active, and methyl tiglate, the least. No mixture of esters was found to be significantly more active than ethyl tiglate alone. In a doseresponse study, bioassay activity increased with dose for both ethyl tiglate andZ10–21. Newly emerged males did not have detectable levels of the esters. All five esters increased as sexual maturity was approached. Ethyl tiglate and ethyl hexanoate were the most abundant in mature males, usually over 15 ng per individual. Ratios among the esters were variable. Male flies also contained an as yet unidentified attractant(s) still more polar than the esters, which was synergistic with the esters and hydrocarbon. Food odors also synergized the synthetic compounds.     

Interruption of response ofDendroctonus brevicomis to attractive pheromone by release of pheromone at several rates and spacings

The number of western pine beetles,Dendroctonus brevicomis, trapped at the center of three 90 × 90-m plots was reduced during a 16-hr period after surrounding an attractive source ofexo-brevicomin, frontalin, and myrcene with 16, 48, or 168 sources of the same attractant. Compounds were released from the center of the plot at 1 mg/16 hr/compound, and from the surrounding sources at 4–280 mg/16 hr/compound. About half of these amounts was released between 1530 hr and 1930 hr, the period of peak beetle flight. No treatment differences were apparent in reduction of catch within the range of release rates and spacings tested. When compounds were released continuously from 168 stations for 17 days, catch at the center of a plot was generally lower than catch before or after this period, but fluctuated daily. More beetles were caught on traps hung on ponderosa pines within a plot and the number of these traps catching beetles was greater when compounds were released from all stations than from only the center station. Beetles caught on traps were attracted into the plots from the surrounding forest, but appeared to be dispersed within the plot when compounds were released from many stations.D. brevicomis attacked at least 91 trees in the plots, of which 25 were killed during two summers.Coleoptera: Scolytidae.Trade names are mentioned solely for information. No endorsement by the U.S. Department of Agriculture is implied.