Identification of the Aggregation Pheromone of the Melon Thrips,Thrips palmi

The objective of this study was to identify the aggregation pheromone of the melon thrips Thrips palmi, a major pest of vegetable and ornamental plants around the world. The species causes damage both through feeding activities and as a vector of tospoviruses, and is a threat to world trade and European horticulture. Improved methods of detecting and controlling this species are needed and the identification of an aggregation pheromone will contribute to this requirement. Bioassays with a Y-tube olfactometer showed that virgin female T. palmi were attracted to the odour of live males, but not to that of live females, and that mixed-age adults of both sexes were attracted to the odour of live males, indicating the presence of a male-produced aggregation pheromone. Examination of the headspace volatiles of adult male T. palmi revealed only one compound that was not found in adult females. It was identified by comparison of its mass spectrum and chromatographic details with those of similar compounds. This compound had a structure like that of the previously identified male-produced aggregation pheromone of the western flower thrips Frankliniella occidentalis. The compound was synthesised and tested in eggplant crops infested with T. palmi in Japan. Significantly greater numbers of both males and females were attracted to traps baited with the putative aggregation pheromone compared to unbaited traps. The aggregation pheromone of T. palmi is thus identified as (R)-lavandulyl 3-methyl-3-butenoate by spectroscopic, chromatographic and behavioural analysis.     

Electroantennogram responses of Apanteles obliquae (Hym., Braconidae) to various infochemicals

Abstract:   The electroantennogram recording technique (EAG) was used to study the olfactory sensitivity of Apanteles obliquae (Hym., Braconidae), a gregarious larval endoparasitoid of Spilosoma obliqua (Walker) (Lep., Arctiidae), to 25 general plant volatiles belonging to alcohol, aldehyde and terpenoid groups and also to volatiles from the host and plant–host complex. The EAG data indicated different olfactory sensitivity between the sexes, not only to individual plant volatiles but also to the volatiles from host and plant–host complex. Females were found to be more responsive than males. However the synthetic sex pheromone blend of the host insect elicited similar EAG responses in both sexes. The EAG data of the present study is correlated with the reported behaviour observed in other parasitoids.     

Use of olfaction for sexual recognition in the subterranean rodent<Emphasis Type="Italic">Ctenomys talarum</Emphasis>

The ability of the tuco-tucoCtenomys talarum Thomas, 1898 to recognize sex by olfactory cues contained in urine, faeces and soiled shavings was tested by using preference tests. Nonbreeding tuco-tucos selected odours from opposite-sex rather than same-sex conspecifics. This pattern differed between sexes: females spent more time sniffing male than female odours for all scent sources whereas males did not show any difference in the time they spent investigating odours of each sex for each tested odour sources. Dissimilarities in odour selections between sexes may be attributed to a different combination of factors involved in olfactory interest for each sex. The function of gender cues recognition is discussed.     

Role of Volatile Semiochemicals in the Host and Mate Location Behavior of Mallodon dasystomus (Coleoptera: Cerambycidae)

Little is known of the role semiochemicals play in the mating systems of longhorned beetles (Coleoptera: Cerambycidae) in the primitive subfamily Prioninae. Mallodon dasystomus (Say), the hardwood stump borer, is a widely distributed prionine native to the southern US. Preferred hosts of M. dasystomus include oak, sweetgum, sugarberry and hackberry; although they also colonize a variety of other hardwoods. Here, we study the mate location behavior of M. dasystomus by testing the hypotheses that the sexes are mutually attracted to volatiles emanating from the larval host and that females release a volatile pheromone that is attractive to males alone. In a Y-tube olfactometer, male and female M. dasystomus responded to volatiles from host material (i.e., sweetgum and sugarberry). However, only males responded to females in the olfactometer, suggesting that females release a volatile sex pheromone. In choice experiments conducted in a greenhouse, we determined that both males and females prefer host over non-host material. In further bioassays in the greenhouse, males chose host material containing a live female over that containing a live male or host material alone. These findings are further evidence of the critical role host volatiles and pheromones play in mating systems of longhorned beetles.     

Olfactometer Responses of Plum Curculio <Emphasis Type="Italic">Conotrachelus nenuphar</Emphasis> (Herbst) (Coleoptera: Curculionidae) to Host Plant Volatiles,Synthetic Grandisoic Acid,and Live Conspecifics

The plum curculio Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae) is a major pest of pome and stone fruit, but will also attack other fruits. Males produce the aggregation pheromone grandisoic acid; emitting only the (+)-enantiomer which is attractive to both sexes of the univoltine and multivoltine strains, while the synthetic racemic mixture contains optical isomers with equal amounts of (+)- and (?)-enantiomers. Synergy between odours can increase trap captures and improve monitoring techniques, therefore tests were performed in a dual-choice olfactometer with odours attractive to plum curculios according to literature to determine 1) under what physiological conditions (mating status, age, starvation period) these odours are attractive or repulsive, 2) if the (+)-enantiomer or the odour of live males synergizes with host plant volatiles, and 3) if there is a difference in response between plum curculio strains. Females were exposed to: benzaldehyde; trans-2-hexenal; apples; extracts of: plums, apples, blueberries; grandisoic acid; and live males. Plum essence was found to be the most attractive host-plant odour for both immature and mature virgin females, and immature whole apples were attractive to starved females, while trans-2-hexenal, McIntosh apple essence, benzaldehyde along with the combination of benzaldehyde and plume essence was found to be repulsive. Starvation, age, and mated status all influence response to odours. No synergistic or additive affects were observed between any of the odour combinations tested, including the combination of both the natural and synthetic pheromone and plum essence or apples.     

多种植物挥发物及马铃薯甲虫聚集素对马铃薯甲虫的引诱作用

马铃薯甲虫Leptinotarsa decemlineata (Say)是马铃薯上一种最主要的毁灭性害虫。由于其严重的抗药性, 急需寻找一种替代杀虫剂的防治方法。本实验从开发马铃薯甲虫引诱剂的目的出发,用“Y”型嗅觉仪测定了马铃薯甲虫对来自植物的7种挥发物单体、8个挥发物混合物配方以及马铃薯甲虫聚集素的行为反应,并进一步进行了田间诱集试验。室内生测结果表明,2-苯乙醇对马铃薯甲虫雌雄虫都有明显的引诱作用。芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯的混合物对马铃薯甲虫雄虫有很强的引诱作用(81.67%), 但是对雌虫引诱作用不明显(63.33%)。在其中加入马铃薯甲虫聚集素不但提高了对雄虫的引诱率（88.33%）,还消除了之前的雌雄性别间反应差异现象,使对雌虫的引诱率达到了83.33%。田间诱集结果显示,芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯+马铃薯甲虫聚集素在所研究的引诱剂配方中引诱效果最好。从而为马铃薯甲虫发生的预测预报和诱杀技术提供了研究基础。     

Evidence for a male-produced aggregation pheromone in Scyphophorus acupunctatus Gyllenhal (Coleoptera: Curculionidae)

Olfactory response of male and female Scyphophorus acupunctatus Gyllenhal (Coleoptera: Curculionidae) to volatiles released from the same or opposite conspecifics alone, or combined with host plant volatiles, was evaluated in the laboratory and field. We also evaluated the response to synthetic Rhynchophorinae pheromones in the laboratory. In laboratory tests, attraction of males and females in Y-tube olfactometer to conspecific males was greater than to females and clean air. Males and females preferred the combination males + agave over agave alone. Both sexes were significantly attracted to 2-methyl-4-heptanol and 2-methyl-4-octanol compared with hexane control. In field trials, weevils were successfully caught in the traps baited with conspecifics and plant material. These field results support those of the laboratory bioassays, showing that males attracted conspecific males and females and addition of plant material enhanced the attraction. These results further suggest that S. acupunctatus produces an aggregation pheromone.     

Concurrent modulation of neuronal and behavioural olfactory responses to sex and host plant cues in a male moth

Mating has profound effects on animal physiology and behaviour, not only in females but also in males, which we show here for olfactory responses. In cotton leafworm moths, Spodoptera littoralis, odour-mediated attraction to sex pheromone and plant volatiles are modulated after mating, producing a behavioural response that matches the physiological condition of the male insect. Unmated males are attracted by upwind flight to sex pheromone released by calling females, as well as to volatiles of lilac flowers and green leaves of the host plant cotton, signalling adult food and mating sites, respectively. Mating temporarily abolishes male attraction to females and host plant odour, but does not diminish attraction to flowers. This behavioural modulation is correlated with a response modulation in the olfactory system, as shown by electro-physiological recordings from antennae and by functional imaging of the antennal lobe, using natural odours and synthetic compounds. An effect of mating on the olfactory responses to pheromone and cotton plant volatiles but not to lilac flowers indicates the presence of functionally independent neural circuits within the olfactory system. Our results indicate that these circuits interconnect and weigh perception of social and habitat odour signals to generate appropriate behavioural responses according to mating state.     

Plant volatiles moderate response to aggregation pheromone in Colorado potato beetle

Abstract:  The orientation of the Colorado potato beetle, Leptinotarsa decemlineata , to a male-produced aggregation pheromone, ( S )-3,7-dimethyl-2-oxo-oct-6-ene-1,3-diol, a three-component plant attractant blend [comprised of ( Z )-3-hexenyl acetate + (±)-linalool + methyl salicylate], and other potato volatiles (nonanal and 2-phenylethanol) were tested. All compounds were previously shown to be active in coupled gas chromatography/electroantennogram experiments. Both the three-component plant attractant blend and 2-phenylethanol were attractive to adult beetles. While male beetles oriented preferentially to both plant attractants vs. a control, females showed little preference. Combining the plant attractants with the pheromone resulted in sexually dimorphic responses similar to those seen with either plant attractant alone. Addition of nonanal abolished the sexually dimorphic response to the pheromone + 2-phenylethanol blend; the new three-component blend was attractive to both sexes. In both laboratory bioassays and field experiments, a combination of the pheromone + the three-component plant attractant was preferred over the plant attractant alone. Thus, it seems likely that combinations of pheromone + plant volatiles may be the most efficacious for field use.     

Specificity of sex pheromones, the role of host plant odour in the olfactory attraction of males, and mate recognition in the aphid Cryptomyzus

Abstract. Choice tests in an olfactometer showed that males of the aphid Cryptomyzus galeopsidis spent more time in the odour of conspecific females than of a sibling species or a more distantly related Cryptomyzus species, which all co-occur on red currant. Analysis of the pheromone-emissions of females showed the presence of a nepetalactol and a nepetalactone, two known aphid sex pheromone components. However, no interspecific difference in the ratio of the components ( c. 30:1) was found, and this ratio did not change as the females aged. Males were only slightly attracted by a synthetic 30:1 mixture of these two components, and in a choice test preferreď the odour of conspecific females. The sex pheromone of Cryptomyzus spp. must thus consist of more than two components, the first time this has been reported for aphids. Host plant odour did not attract males or enhance the activity of the pheromone.     

Aggregation pheromone in Sternechus subsignatus (Coleoptera: Curculionidae): olfactory behaviour and temporal pattern of emission

The Brazilian soybean stalk weevil, Sternechus subsignatus Boehman (Coleoptera: Curculionidae), is a pest of economic importance in many regions of Brazil. Volatiles from both sexes of S. subsignatus were collected by aeration and the behavioural response of males and females was evaluated using a Y‐olfactometer. The results obtained demonstrate that the communication in S. subsignatus is mediated by aggregation pheromone as both sexes were attracted to host plant (HP) volatiles, and this attraction was increased by the addition of male volatiles. At least five male‐specific compounds (1–5) were detected in the chromatographic analysis, providing chemical support to the behavioural data. Release of these volatiles is dependent on the presence of the HP as the amount of compounds differs significantly when volatiles are collected from weevils with or without access to food. The release takes place mainly during photophase, showing a peak between 4 and 6 h after its beginning, which is also a peak of the insect activity in the field. Studies are underway to elucidate the structures of these putative aggregation pheromone components, in order to evaluate their biological activity under laboratory and field conditions.     

Synergism of pheromone and host‐plant volatile blends in the attraction of Grapholita molesta males

Control of Grapholita molesta (Busck) (Lepidoptera: Tortricidae), a major pest of stone and pome fruits, is successfully achieved by mating disruption. Under these conditions, tools other than conventional pheromone dispensers are needed for flight monitoring. The objective of the present work was to determine whether plant volatiles synergize male G. molesta attraction to a suboptimal dose of synthetic sex pheromone. The plant blend (referred to as 5VB), a mixture of three green leaf volatiles [(Z)‐3‐hexenyl acetate, (Z)‐3‐hexenol, and (E)‐2‐hexenal] and two aromatics [benzaldehyde (BZA) and benzonitrile (BZN)], was added to the suboptimal pheromone dose (2 ng on filter paper) in log steps (up to 10 000× the pheromone dose) to test synergism of pheromone and plant blends. In addition, the effect of individual plant volatiles on male responses was investigated by adding to the suboptimal pheromone dose each of the four‐compound plant‐volatile blends, resulting from eliminating one volatile from the 5VB at a time, or each plant volatile alone. Flight behaviour and the time to reach the source were recorded. The 5VB alone was not attractive to G. molesta males, but at a ratio of 1:1 000 (Ph:5VB) or higher, the attractiveness of the suboptimal pheromone dose increased, to a level similar to that of the optimal pheromone dose (10 ng). All tested plant volatiles, except BZA, synergized the response to the pheromone when added individually, but only (Z)‐3‐hexenol and BZN did so to a level not significantly different from the Ph:5VB blend. Aromatics had a stronger effect than green leaf volatiles (GLVs), because their removal, but not the removal of GLVs, decreased landing responses. The addition of the 5VB decreased significantly the time males needed to reach the odour source. The observed enhanced male attraction to mixtures of pheromone and plant volatiles will facilitate the development of lures for G. molesta adult flight monitoring.     

Electroantennogram responses of the carrot fly, Psila rosae, to volatile plant components

ABSTRACT. Electroantennogram (EAG) responses of male and female carrot flies, Psila rosae F. (Diptera: Psilidae), were recorded to thirty-six volatile plant constituents. The most distinct EAG responses were obtained to: (1) the general green leaf volatiles 1-hexanol, trans-2-hexen-1-ol and cis-3-hexen-1-ol, their isomers cis-2-hexen-1-ol and trans-3-hexen-1-ol, the alcohol 1-heptanol, the ester cis-3-hexenyl acetate and the leaf aldehydes hexanal and trans-2-hexenal, and (2) from four compounds associated with the umbelliferous host plants of this insect, namely trans-methyl-iso-eugenol, β-caryophyllene, linalool and trans-2-nonenal. Higher responses were elicited by the leaf aldehydes than by the corresponding alcohols. Although the absolute amplitude of the female response was over twice that of the male, there were no differences between the relative responses to the compounds tested in both sexes, with the exception of a much higher response to the leaf aldehydes in the male. The shape of the EAG evoked by the various compounds was consistently different, with the slowest recovery being recorded for trans-methyl-iso-eugenol. While the antennal olfactory receptors of the carrot fly are sensitive to the closely related general green leaf volatiles, they are most specifically tuned to the aldehyde component of this green odour complex. In addition, the ability of this insect to discriminate between different plants may be augmented by the perception of a group of more host specific volatiles. The conformity of the responses of males and females to the compounds tested may indicate that host plant volatiles plays an additional role as an aggregation cue for both sexes.     

Optimizing a pheromone lure for the sugar beet root maggot fly,Tetanops myopaeformis

We previously described a putative aggregation pheromone in adults of the sugar beet root maggot, Tetanops myopaeformis (von Röder) (Diptera: Ulidiidae), comprising nine compounds identified from males. Here, we conducted a series of experiments aimed at simplifying the blend of compounds necessary to achieve attraction as well as determining the dose that maximizes captures when formulated into an attractant lure. In all experiments, females showed stronger and more consistent evidence of attraction than males. White sticky traps baited with different blends of pheromone compounds that included the major component, (R)-(−)-2-nonanol, showed significantly higher female captures relative to those baited with blends that excluded the major component. (R)-(−)-2-nonanol alone was at least as effective as any blend that included this compound with other minor pheromone components. Lures using racemic 2-nonanol were as effective as the (R) enantiomer for both females and males, with some evidence of weak attraction to the (S) enantiomer (which is not produced by males) observed as well. Maximum capture rates using racemic 2-nonanol were estimated to occur with doses of ca. 795.5 and 621.6 mg for females and males, respectively. Addition of 2-nonanol lures to standard orange sticky stake traps currently used to monitor flies increased captures of both sexes. The pheromone lure developed here could improve trapping efficiency of current monitoring programs for T. myopaeformis and may also be used to develop other management tools for this important pest of sugar beet.     

Sex‐specific responses of Asian citrus psyllid to volatiles of conspecific and host‐plant origin

Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is the primary vector of Candidatus Liberibacter spp. bacteria that cause citrus greening, a disease of worldwide importance. Olfactometry was employed to test responses of D. citri to odours from intact citrus plants (Mexican lime, Citrus aurantifolia, sour orange, Citrus aurantium, Marsh grapefruit, Citrus paradisi and Valencia orange, Citrus sinensis), citrus plants previously infested with D. citri, and odours of conspecifics including nymphs, adult insects of same and opposite sex, and their products (honeydew), both alone and in combination. In contrast to other studies, psyllids of both sexes were attracted to volatiles of undamaged Mexican lime leaves, whereas undamaged grapefruit attracted only females, and leaves of Valencia and sour orange did not attract either sex. All four plant species attracted female psyllids when previously infested, but only Mexican lime and sour orange‐attracted males. Thus, Citrus species appear to vary in the production of both constituitive and induced volatiles that attract adult psyllids. Volatiles emitted by nymphs did not attract either sex, but psyllid honeydew was attractive to males, likely due to female pheromone residues. Males oriented to the odour of females, whereas the reverse was not true, and neither males nor females oriented to same‐sex volatiles. The addition of conspecific cues (adults, nymphs or honeydew) did not increase female attraction to previously infested leaves, but male response was increased by the presence of adults and honeydew, regardless of plant species. Thus, female psyllids appear to orient more strongly to volatiles of plant origin, whereas males respond more strongly to cues emanating from females and conspecific excretions. These results suggest that female psyllids drive the initial colonization of host plants, whereas males orient to females and infested plants. Identification of the specific volatiles involved may permit their use in monitoring and management of this pest.     

The Role of Chemical Cues in Host and Mate Location in the Pear Psylla Cacopsylla bidens (Homoptera: Psyllidae)

Chemical communication was shown to play a role in the pear psylla, Cacopsylla bidens. Electrophysiological (EAG) and behavioral responses were investigated in males and females pear psylla . Males were found to be attracted to females, and especially to those on host plants, but not to males, uninfested host plants, or plants infested with conspecific larvae. On the other hand, females were not attracted to males or females but displayed some attraction to host plants. Furthermore, females showed a preference for uninfested pear versus plants infested with conspecific larvae. The antennae of males gave highest electroantenographic response to volatiles from pears infested with females but not males, while females, responded also toward the volatiles of pear alone. These results indicate that females of C. bidens emit sex pheromones that are attractive to the males and suggest that, host volatiles may play a role in host selection by pear psylla females.     

A six-arm olfactometer permitting simultaneous observation of insect attraction and odour trapping

Abstract.  Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose–response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed.     

Sex pheromone components of female Cornutiplusia circumflexa

Capillary gas chromatography indicated the presence of (Z)-7-dodecenyl alcohol and (Z)-7-dodecenyl acetate in the abdominal tip extracts of female Cornutiplusia circumflexa (L.) (Lepidoptera: Noctuidae: Plusiinae) moths. Gas chromatography combined with mass spectrometry and dimethyl disulfide derivatization confirmed the structure of these two principal pheromone components. In a flight-tunnel bioassay, a 5:1 blend of alcohol and acetate elicited the complete courtship sequence by males. The response was comparable to that evoked by a live virgin female, but males spent significantly longer time at calling females than at the synthetic source. The 5:1 blend was also attractive to male C. circumflexa in the field, as indicated by catches in traps baited with this mixture. (Z)-7-dodecenyl alcohol alone was slightly attractive whereas (Z)-7-dodecenyl acetate alone was completely inactive. C. circumflexa is the first reported Plusiinae species which utilizes (Z)-7-dodecenyl alcohol as the major pheromone component.     

Host plant volatiles induce oriented flight behaviour in male European grapevine moths, Lobesia botrana

The European grapevine moth Lobesia botrana relies on a female produced sex pheromone for long-distance mate finding. Grapevine moth males compete heavily during limited time windows for females. The aim of this study was to investigate the perception of host plant volatiles by grapevine moth males and whether such compounds elicit upwind oriented flights. We compared five host plant headspace extracts by means of gas chromatography linked electroantennogram (EAG) recording. We identified 12 common host plant volatiles (aliphatic esters, aldehydes, and alcohols, aromatic compounds and terpenes) that elicit EAG responses from grapevine moth males and that occur in at least three of the host plant volatile headspace extracts tested. Subsequently the behavioural response of grapevine moth males to four these compounds presented singly and in mixtures (1-hexanol, 1-octen-3-ol, (Z)-3-hexenyl acetate and (E)-β-caryophyllene) was recorded in a wind tunnel. Grapevine moth males engaged in upwind flights to all of four compounds when released singly at 10,000 pg/min and to all, except 1-octen-3-ol, when released at 100 pg/min. A blend of the four host plant volatiles released at 10,000 pg/min and mixed at a ratio based on the analysis of Vitis vinifera cv. Solaris volatile emissions attracted significantly more males than any single compound. Grapevine moth males perceive and respond to host plant volatiles at biologically relevant levels indicating that host plant volatiles figure as olfactory cues and that L. botrana males can discern places where the likelihood of encountering females is higher.     

蚜虫雄蚜与雌性母对性信息素及植物挥发物的田间反应

田间观察了桃蚜Myzus persicae (Sulzer)、绣线菊蚜Aphis spiraecola Patch 、山楂圆疣蚜Ovatus crataegarius (Walker)等3种蚜虫对性信息素[(4aS,7S,7aR)-荆芥内酯和 (1R,4aS,7S,7aR)-荆芥醇]的反应,并且调查了性信息素与植物挥发物对桃蚜的田间引诱活性的相互作用.在有冬寄主或夏寄主植物的田中,性信息素诱捕器诱捕到桃蚜雄蚜与雌性母的数量显著多于对照诱捕器的诱捕数;但在非寄主植物的田中,却引诱不到桃蚜.苯甲醛(冬寄主植物桃树Prunus persica的主要挥发物组分之一)能够增强桃蚜雄蚜的引诱作用.绣线菊蚜雄蚜和雌性母对植物中提取的荆芥内酯有反应,而山楂圆疣蚜雄蚜和雌性母对植物中提取的和人工合成的荆芥内酯都没有反应,但对荆芥醇有反应.并且在荆芥醇中添加荆芥内酯之后对山楂圆疣蚜雄蚜引诱活性显著提高.还讨论了雌性蚜产生化合物被雄蚜作为性信息素、被雌性母作为聚集信息素以及性信息素与寄主植物挥发物的相互作用.