Fine-scale resolution of closely spaced pheromone and antagonist filaments by flying male Helicoverpa zea

The limits of a male moth's ability to resolve closely spaced odor filaments have been investigated. Male Helicoverpa zea normally respond to their conspecific sex pheromone blend by exhibiting an upwind flight, which culminates in source contact by at least 50% of the bioassayed individuals. When loaded onto the same filter paper source containing this hitherto attractive pheromone blend, or onto a separate filter paper and co-emitted from the same pipette source with pheromone, (Z)-11-hexadecenyl acetate severely reduced upwind flight and source contact by male H. zea. A similar level of upwind flight inhibition was recorded when the antagonist (Z)-11-hexadecenyl acetate was emitted from its own point source placed 1 mm upwind of the pheromone point source, both plumes being simultaneously emitted in a continuous mode to form a confluent strand. However, (Z)-11-hexadecenyl acetate was less effective in reducing upwind flight and source contact when it was isolated and pulsed from its own source, placed 1 mm either upwind, downwind or cross-wind of a pipette source from which pheromone was simultaneously being pulsed, such that both filaments were separated in time by 0.001–0. 003 s. These results suggest that male H. zea are able to distinguish between odor sources separated by as little as 1 mm in space and 0.001 s in time. Accepted: 31 March 1999     

Helicoverpa zea males (Lepidoptera: Noctuidae) respond to the intermittent fine structure of their sex pheromone plume and an antagonist in a flight tunnel

Abstract. We investigated the behavioural response of male Helicoverpa zea (Boddie) to the fine-scale structure of an odour plume experimentally modified in a wind tunnel by using an air-pulsing device. Male H. zea flew upwind to pulsed filaments of a binary pheromone blend of (Z)-11-hexadecanal (Z11-16:Ald) and (Z)-9-hexadecanal (Z9-16:Ald) in the ratio of 20:1. Sustained upwind flight in experimentally altered intermittent plumes was dependent on concentration, as well as the frequency of generation of odour filaments. At a loading of 10μg of the major pheromone component, Zll-16:Ald, which gave an emission rate of approximately that released by a female H. zea , sustained upwind flight and source contact correlated positively with filament delivery rate, becoming significant at a minimum filament delivery rate of 2/s. Decreases in upwind progress and source location were recorded at a loading of 1 μg of Z11-16:Ald. At this suboptimal dosage, a high filament generation rate of 10/s was necessary for significant upwind progress and source contact. When an interspecific compound: (Z)-11-hexadecenyl acetate (Z11-16:OAc), was added to the attractive pheromone binary aldehyde blend of H. zea at a proportion of 10% of the major pheromone component, and pulsed from the same source, there was a significant reduction in sustained upwind progress and source location by males, indicating that Z11-16:OAc is antagonistic to the upwind progress of H. zea. However, Z11-16:OAc was less antagonistic when its filaments were isolated and alternated with pheromone filaments, indicating a strong effect of the synchronous arrival of odour filaments on the antenna needed for antagonism of upwind flight.     

棉铃虫二种性诱芯田间诱蛾效果比较

棉铃虫Helicoverpa armigera(Hübner)是我国最严重害虫之一。为了提高监测和诱捕效率,2009年在山西和新疆进行2种性诱芯诱捕棉铃虫的田间试验,2种性诱芯分别来自中国科学院动物研究所和北京中捷四方公司,2种诱芯处理分别重复2次和6次,诱捕器分别为水盆和笼罩。试验结果:在山西试点,动物所诱芯的诱蛾数是中捷四方公司的8.60倍;在新疆试点,则达到11.89倍。以上差异均达极显著水准(P<0.01)。同时,在山西试点,动物所诱芯的有效期长达62d,比中捷四方公司的长14d。据此认为动物所棉铃虫性诱芯值得应用推广。     

Antennal lobe projection destinations of Helicoverpa zea male olfactory receptor neurons responsive to heliothine sex pheromone components

We used single sensillum recordings to define male Helicoverpa zea olfactory receptor neuron physiology followed by cobalt staining to trace the axons to destination glomeruli of the antennal lobe. Receptor neurons in type A sensilla that respond to the major pheromone component, (Z)-11-hexadecenal, projected axons to the cumulus of the macroglomerular complex (MGC). In approximately 40% of these sensilla a second receptor neuron was stained that projected consistently to a specific glomerulus residing in a previously unrecognized glomerular complex with six other glomeruli stationed immediately posterior to the MGC. Cobalt staining corroborated by calcium imaging showed that receptor neurons in type C sensilla sensitive to (Z)-9-hexadecenal projected to the dorsomedial posterior glomerulus of the MGC, whereas the co-compartmentalized antagonist-sensitive neurons projected to the dorsomedial anterior glomerulus. We also discovered that the olfactory receptor neurons in type B sensilla exhibit the same axonal projections as those in type C sensilla. Thus, it seems that type B sensilla are anatomically type C with regard to the projection destinations of the two receptor neurons, but physiologically one of the receptor neurons is now unresponsive to everything except (Z)-9-tetradecenal, and the other responds to none of the pheromone-related odorants tested.     

Neurons discovered in male Helicoverpa zea antennae that correlate with pheromone-mediated attraction and interspecific antagonism

Responses of single receptor neurons in the antennae of male Helicoverpa zea to sex pheromone components and to behavioral antagonists were recorded using a cut-sensillum extracellular recording technique. Three types of sensilla were identified from sampling 325 male-specific sensilla trichodea located at the lateral edge of antennomeres. The majority of these sensilla (71%) contained a receptor neuron tuned to the principal sex pheromone component (Z)-11-hexadecenal. A second sensillar type (10%) contained a receptor neuron that responded only to (Z)-9-tetradecenal. A third sensillar type (19%) contained a large-spiking neuron tuned to the secondary pheromone component (Z)-9-hexadecenal, but this neuron also could be stimulated to equivalent spike frequencies by the same emitted amounts of (Z)-9-tetradecenal. A smaller-spiking neuron in this sensillar type responded to two compounds known to act only as behavioral antagonists, (Z)-11-hexadecen-1-ol and (Z)-11-hexadecenyl acetate, and to (Z)-9-tetradecenal. Cross-adaptation studies confirmed the presence of one large- and one small-spiking neuron in the third sensillar type. Dose-response studies correlated to collected stimuli amounts showed that the large-spiking neuron in the third sensillar type was equally tuned to (Z)-9-hexadecenal and (Z)-9-tetradecenal, whereas the smaller-spiking neuron was far more sensitive to (Z)-11-hexadecen-1-ol and to (Z)-11-hexadecenyl acetate than to (Z)-9-tetradecenal. Accepted: 29 September 1997     

Structural organization of the sex pheromone gland in Helicoverpa zea in relation to pheromone production and release

Morphological location of the sex pheromone producing area in the ovipositor of the female corn earworm Helicoverpa zea, was correlated with gas chromatographic analysis of the extracted pheromone. Histological studies showed that the pheromone gland occupied an almost complete ring of specialized columnar cells between the 8th and 9th abdominal segments. Ultrastructure of the pheromone gland cells revealed distinct features such as microvilli, pockets of granular material, intercellular canals with abundant desmosomes. Apparent changes in some of these features are associated with phases of pheromone production and non-production. Examination of the tissue with low temperature scanning electron microscopy showed the presence of excreted droplets at the tips of cuticular hairs in the glandular area during the period of pheromone production.     

Scoliopteryx libatrix (L.) (Lep., Noctuidae) male reaction to the synthetic main sex pheromone component and its isomers – wind tunnel and field investigations

Abstract: The four stereoisomers of 6,13-methylheneicosene were tested with Scoliopteryx libatrix (L.) males in a wind tunnel. The most active isomer was (Z6,S13)-methylheneicosene which had been identified as the main sex pheromone component of the species. Some medium activity was found for (Z6,R13)- and (E6,S13)-, while the (E6,R13)-stereoisomer showed the lowest activity. These results were confirmed in field tests with sticky traps. Monitoring of S. libatrix at two sites showed that the flight of the insects started at the end of March to the beginning of April and lasted till August to the beginning of September. It was concluded that the males of the second generation were not attracted by the pheromone.     

Antagonistic Effect of (Z)-11-Hexadecen-1-ol on the Pheromone-Mediated Flight of Helicoverpa zea (Boddie) (Lepidoptera:Noctuidae)

Flight-tunnel experiments were conducted using Helicoverpa zea males to determine whether or not (Z)-11-hexadecen-1-ol (Z11-16:OH), a compound emitted by another heliothine moth species, Heliothis subflexa, is a behavioral antagonist when admixed with the two-component pheromone blend of H. zea. Males were less likely to fly upwind all the way to the source when 0.3% Z11-16:OH was present in the blend. Even 0.1% Z11-16:OH caused differences in the flight behavior of H. zea males; they steered more off the windline than males responding to the pheromone blend alone, resulting in more oblique track angles. Thus Z11-16:OH appears to act antagonistically, along with another compound, (Z)-11-hexadecen-1-ol acetate (Z11-16:Ac), when it is added to the H. zea pheromone blend.     

蛾类昆虫雄性信息素及其功能

昆虫性信息素是两性通讯系统的基础,其中雄性信息素的研究相对较少。本文综述了蛾类昆虫雄性信息素的研究进展。迄今已鉴定出40余种蛾类昆虫的雄性信息素,其行为学功能主要有对雌性的引诱和激欲、对同种雄性的抑制及种间隔离等。     

Temperature modulation of behavioural thresholds controlling male moth sex pheromone response specificity

ABSTRACT. The response specificity of male Oriental fruit moths, Grapholita molesta (Busck) and pink bollworm moths, Pectinophora gossypiella (Saunders), to different blends and doses of pheromone is altered dramatically by temperature. When acclimated and tested in a flight tunnel at 20^oC males of both species exhibit a high degree of specificity, with peak response levels occurring to a narrow range of blend-dose combinations close to the natural blend. When tested at 26^oC, however, males exhibit a significantly lower degree of specificity, with peak response occurring to a broader range of treatments. The change in response specificity results from shifts in behavioural threshold effects influencing plume orientation and initiation of upwind flight, as well as from arrestment of upwind flight, occurring later in the flight sequence. The observed changes in male behaviour suggest that the effect of temperature is directly on neural pathways involved in the perception of odour, and not simply the result of an increase in motor activity or a significant change in the release rate of the pheromone. The results support the threshold hypothesis for pheromone perception (Roelofs, 1978) as a general principle in the Lepidoptera, but also show that the degree of response specificity can be significantly affected by temperature.     

Responses of male California red scale to sex pheromone isomers

The California red scale, Aonidiella aurantii (Maskell), sex pheromone components were previously identified as two independently attractive norsesquiterpenes. The four possible optical isomers of one and the four possible geometric and optical isomers of the other were synthesized and tested for male California red scale attractiveness in field tests and greenhouse bioassays. The results showed that there was enantiomeric and geometric specificity and only 1 isomer of each component was significantly more active than the others. The active isomers were (3S,6R)-3-methyl-6-isopropenyl-9 decen-1-yl acetate and (3Z,6R)-3-methyl-6-isopropenyl-3,9-decadien-1-yl acetate. The presence of other isomers, including the synthetic analogue 3-methyl-6-isopropylidene-9-decen-1-yl acetate, had no effect on trap catch.     

Circadian rhythmicity and development of the behavioural response to sex pheromone in male brown-banded cockroaches, Supella longipalpa

Abstract The effects of age, time of day and pheromone concentration on the responsiveness of the male brown-banded cockroach, Supella longipalpa (F.), to female volatile sex pheromone were examined. Male response increased with age and peaked 10 days after the imaginal moult. A diel periodicity of responsiveness was observed, with greater responses in the scotophase. The behavioural response was dose-dependent and its diel periodicity was most obvious when intermediate pheromone concentrations (e.g. 0.01 female equivalents) were used. Higher concentrations obscured the periodicity by eliciting greater responses in the photophase, while the lower concentrations did so by decreasing the response in the scotophase. Following entrainment to a LD 12:12 h cycle, the periodicity of response to 0.01 female equivalents of pheromone persisted for at least 54 h under continuous darkness, further demonstrating a true circadian rhythm.     

雄性昆虫性信息素的研究进展

全面概述雄性昆虫性信息素的研究进展。主要从性信息素的释放机制及其作用、已发现的雄性昆虫性信息素及其化学结构进行阐述。通过不同部位释放的雄性昆虫性信息素种类是多样的,但其化学结构的差异又是细微的,雄性昆虫则利用这些差异来引诱雌性昆虫、促进交配和生殖隔离等。最后对雄性昆虫性信息素的研究作了展望。     

A direct search approach to characterize the sex pheromone composition giving maximal male response

ABSTRACT. Response of male moths to sex pheromones differs markedly depending on blend composition. Natural selection can therefore be expected to favour males responding maximally to the blend produced by most females in the population. Thus it is relevant to quantify male response by the blend to which the males of a population respond maximally.
To do this, wind tunnel experiments were carried out where males were observed while responding to blends of synthetic compounds emitted from rubber septa. By stepwise improvement it was shown that the blend to which Yponomeuta cagnagellus responds maximally comprises (Z)11-tetradecenyl acetate (Z11-14Ac), ( E )11-tetradecenyl acetate (E11-14Ac), (Z)11-tetradecenol (Z11-140H), and tetradecyl acetate (14Ac) in the ratio 100(=50μg):1.5:20:45.
The innovate aspects of the method used to obtain this result, are the use of a direct search approach as an integral part of the experimental procedure and the use of a summary variable to characterize male response. These aspects are discussed using the data on the response of Y. cagnagellus males.     

Responses to aphid sex pheromones by the pea aphid parasitoids Aphidius ervi and Aphidius eadyi

Aphidius ervi and Aphidius eadyi, two parasitoids of the pea aphid Acyrthosiphon pisum, were attracted to components of the aphid sex pheromone in laboratory bioassays. Pre-test experience with host aphids in the presence of aphid sex pheromone did not affect the response of A. ervi to pheromone in a 4-way olfactometer, compared with that of naive parasitoids. Aphidius ervi females exposed only to the pheromone prior to testing did not respond in the olfactometer, suggesting habituation to the foraging cue by the parasitoid. In a wind tunnel, aphid sex pheromone increased the attraction of A. ervi to the plant-host complex (Vicia faba/A. pisum), suggesting an additive effect when two different foraging cues are present simultaneously.     

Disruption of male spruce budworm orientation to female moths by pheromone and pheromone analogues in a wind tunnel

Abstract. The ability of male spruce budworm moths, Choristoneura fumiferana (Clem.), to orientate to and locate virgin female moths was investigated in a laboratory wind tunnel. When presented with two caged ‘target female’ moths, more than 99% of the male moths locked on and flew directly to the females. When confined for 3 h before release, downwind of a three-dimensional array of thirty-five evenly distributed female moths (a density of 40/m³), over 90% of the males still successfully oriented to one or another of the females, although not always to the target females. With an array of thirty-five septa containing either E-11-tetradecenyl acetate on E-11-tetradecenol there was no visible change in behaviour; >90% of the males flew directly to the two target females, which indicates that neither the acetate nor the alcohol has potential for mating disruption. An array of thirty-five septa containing a 955 blend of E- and Z-11-tetradecenal, the main components of the spruce budworm sex pheromone, significantly reduced the numbers of male moths able to locate the target females, and effects were concentration-dependent. The highest concentration, which gave a release rate similar to that of a calling female, produced the same level of disruption as the thirty-five females, but sensory fatigue became a factor in addition to ‘false trail following’. Lower concentrations were significantly less effective, with no evidence of sensory fatigue. At all concentrations significant numbers of males flew to the septa (false trail following), but when male moths were left uncaged in the tunnel for a further 3h, most eventually located the females. This suggests that false trail following alone will not prevent mating. The level of disruption was not improved by the addition of 2% tetradecanal to the 955 blend. Similar levels of disruption to the 955 blend were obtained with an 80:20 blend. A 5050 blend still caused a significant amount of false trail following, but less than the other blends. The ability of male spruce budworm to orientate to off-blend trails was enhanced if the males were first exposed to the natural, female-produced pheromone.     

Characteristics of 13 polymorphic microsatellite markers in the corn earworm,Helicoverpa zea (Lepidoptera: Noctuidae)

Thirteen polymorphic microsatellite loci suitable for population genetic studies of Helicoverpa zea were discovered by screening partial genomic libraries enriched for microsatellite sequences. Insects collected (N = 96) in Stoneville, Mississippi were used to characterize these markers. The observed and effective number of alleles per locus ranged from two to nine (average 4.46) and from 1.07 to 2.45 (average 1.81), respectively. Fisher exact tests detected significant deviations from Hardy–Weinberg equilibrium at three loci, probably due to inbreeding, null alleles, or Wahlund's effect. Significant genotypic disequilibrium was not detected between any pair of loci.     

棉铃虫成虫对性信息素的电生理和行为反应研究

通过EAG和风洞实验,研究了棉铃虫雌雄成虫对性信息素组分和诱芯(Z-11-16Ald∶Z-9-16Ald=97∶3)的电生理反应。其中棉铃虫雌、雄蛾对诱芯的平均EAG反应测定值分别为1.06mV和4.32mV,分别高出对照(无性信息素空白诱芯)0.67mV和0.366mV,差异均达到极显著水平(雌蛾：t＝25.020, P≤0.01;雄蛾：t＝44.269,P≤0.01);棉铃虫雌蛾对性信息素组分(Z-11-16-Ald和Z-9.16Ald)的EAG反应值随浓度增加而增加;雄蛾在被剪除触角后与雌蛾不能正常交配,而雌蛾在被剪除触角后仍有40%的交配率,比正常雌雄蛾的交配率(70%)有所下降;在风洞实验中,雄蛾没有顺风远离诱源的飞行行为,趋向诱源的比率为81.8%,与对照有显著差异。研究表明性信息素组分对棉铃虫的交配活动有明显的影响。     

Sustained-flight tunnel responses of male lightbrown apple moth to synthetic sex pheromone

ABSTRACT. The upwind flight response of individual male lightbrown apple moths, Epiphyas postvittana (Walker), to twenty combinations of the two pheromone components, (E)-11-tetradecenyl acetate (E11-14:OAc) and (E,E)-9,11-tetradecadienyl acetate (E,E-9,11–14:OAc), was observed in a sustained-flight tunnel. At the lowest dosage of E11-14:OAc tested (3 μg), a significantly greater percentage of males flew upwind to and landed at a source emitting 10% E1,E-9, 11–14:OAc than to all other sources. As the source dosage was increased, males showed decreased specificity of upwind flight to ratios of the two components. However, at the highest dosage of E11-14:OAc tested (300 μg), the response specificity of males shifted to blends containing lower percentages of E,E-9,11–14:OAc. The addition of tetradecyl acetate to a blend of the two components did not result in any detectable increase in male response. Analyses of the pheromone glands of individual female Iightbrown apple moths showed that females produced a range of ratios of E11-14;OAc:E,E-9,11–14:OAc from 100:2.2 to 100:11.4 with a median of approximately 100:7, reasonably paralleling the peak response of males. Pre-exposure of males to a blend of the two components, followed by exposure to E11-14:OAc alone (previously reported as a 'memory' effect) resulted in no significant response to E11-14:OAc alone.     

半翅目昆虫性信息素的研究进展

本文主要从半翅目昆虫性信息素的提取鉴定方法、分泌部位、性信息素成分、在防治中的应用以及存在的问题等几个方面综述了半翅目昆虫性信息素的研究进展。