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Bioactive fractions containing methyl eugenol-derived sex pheromonal components in haemolymph of the male fruit fly Bactrocera dorsalis (Diptera: Tephritidae)

Hee AK, Tan KH

Bull. Entomol. Res., 2005 Dec;95(6):615-20.
PMID: 16336709 DOI: 10.1079/BER2005392

Sex pheromonal components of the tephritid fruit fly Bactrocera dorsalis (Hendel), 2-allyl-4,5-dimethoxyphenol and (E)-coniferyl alcohol, are biosynthesized from a highly potent male attractant, methyl eugenol, then sequestered and stored in the rectal gland prior to their release during courtship at dusk. These sex pheromonal components have been detected in the haemolymph and crop organ. Hence, attempts were made to separate and identify the haemolymph fractions which contained the sex pheromonal components. Identification of these bioactive fractions in methyl eugenol-fed male flies using gel filtration column chromatography and biodetection using live male flies showed two fractions as highly attractive to conspecific males. These fractions show a significant increase in protein absorbance in the elution profile of haemolymph from methyl eugenol-fed males compared with that from methyl eugenol-deprived males. The molecular mass of these bioactive fractions as determined by using gel filtration was in the peptide range of 3.3 to 5.5 kDa. Subsequent gas chromatography-mass spectrometry analyses further confirmed the presence of the pheromonal components in the bioactive fractions. The presence of these methyl eugenol-derived sex pheromonal components in specific haemolymph fractions suggests the involvement of a sex pheromone binding complex.
Male sex pheromonal components derived from methyl eugenol in the hemolymph of the fruit fly Bactrocera papayae

Hee AK, Tan KH

J Chem Ecol, 2004 Nov;30(11):2127-38.
PMID: 15672660 DOI: 10.1023/B:JOEC.0000048778.02561.70

Pharmacophagy of methyl eugenol (ME)--a highly potent male attractant, by Bactrocera papayae results in the hydroxylation of ME to sex pheromonal components, 2-ally-4,5-dimethoxyphenol (DMP) and (E)-coniferyl alcohol (CF). These compounds, which are also male attractants, are then sequestered and stored in the rectal gland prior to their release during courtship at dusk. Chemical analyses of the digestive tract (excluding the crop and rectal gland) showed the absence of the sex pheromonal components and their precursor, ME. However, B. papayae males were attracted to and fed on the ME-fed male hemolymph extracts but not on hemolymph extracts of ME-deprived males. After thin layer chromatography in a hexane:ethyl acetate solvent system, flies were attracted to and fed on the original point on the TLC plate where the hemolymph extract had been spotted, suggesting that the pheromone components were bound in polar complexes. Chemical analyses of the ME-fed male hemolymph and crop extracts revealed the presence of the sex pheromonal components. The presence of the ME-derived pheromonal components and the absence of ME in the hemolymph suggest that the hemolymph is involved in the transportation of sex pheromonal components from the crop to the rectal gland.
Fulltext Recent Advancements in Studies on Chemosensory Mechanisms Underlying Detection of Semiochemicals in Dacini Fruit Flies of Economic Importance (Diptera: Tephritidae)

Ono H, Hee AK, Jiang H

Insects, 2021 Jan 26;12(2).
PMID: 33530622 DOI: 10.3390/insects12020106

Dacini fruit flies mainly contain two genera, Bactrocera and Zeugodacus, and include many important pests of fruits and vegetables. Their life cycle is affected by various environmental cues. Among them, multiple characteristic semiochemicals have remarkable effects on their reproductive and host-finding behaviors. Notably, floral fragrances released from so-called fruit fly orchids strongly attract males of several Dacini fruit fly species. Focusing on the strong attraction of male flies to particular chemicals, natural and synthetic lures have been used for pest management. Thus, the perception of semiochemicals is important to understand environmental adaptation in Dacini fruit flies. Since next-generation sequencers are available, a large number of chemosensory-related genes have been identified in Dacini fruit flies, as well as other insects. Furthermore, recent studies have succeeded in the functional analyses of olfactory receptors in response to semiochemicals. Thus, characterization of molecular components required for chemoreception is under way. However, the mechanisms underlying chemoreception remain largely unknown. This paper reviews recent findings on peripheral mechanisms in the perception of odors in Dacini fruit flies, describing related studies in other dipteran species, mainly the model insect Drosophilamelanogaster. Based on the review, important themes for future research have also been discussed.
Fulltext Comparative sensitivity to methyl eugenol of four putative Bactrocera dorsalis complex sibling species - further evidence that they belong to one and the same species B. dorsalis

Hee AK, Ooi YS, Wee SL, Tan KH

Zookeys, 2015.
PMID: 26798265 DOI: 10.3897/zookeys.540.6099

Males of certain species belonging to the Bactrocera dorsalis complex are strongly attracted to, and readily feed on methyl eugenol (ME), a plant secondary compound that is found in over 480 plant species worldwide. Amongst those species is one of the world's most severe fruit pests the Oriental fruit fly, Bactrocera dorsalis s.s., and the former taxonomic species Bactrocera invadens, Bactrocera papayae and Bactrocera philippinensis. The latter species have been recently synonymised with Bactrocera dorsalis based on their very similar morphology, mating compatibility, molecular genetics and identical sex pheromones following consumption of ME. Previous studies have shown that male fruit fly responsiveness to lures is a unique phenomenon that is dose species-specific, besides showing a close correlation to sexual maturity attainment. This led us to use ME sensitivity as a behavioural parameter to test if Bactrocera dorsalis and the three former taxonomic species had similar sensitivity towards odours of ME. Using Probit analysis, we estimated the median dose of ME required to elicit species' positive response in 50% of each population tested (ED50). ED50 values were compared between Bactrocera dorsalis and the former species. Our results showed no significant differences between Bactrocera dorsalis s.s., and the former Bactrocera invadens, Bactrocera papayae and Bactrocera philippinensis in their response to ME. We consider that the Bactrocera males' sensitivity to ME may be a useful behavioural parameter for species delimitation and, in addition to other integrative taxonomic tools used, provides further supportive evidence that the four taxa belong to one and the same biological species, Bactrocera dorsalis.
Fulltext Involvement of the Antennal and Maxillary Palp Structures in Detection and Response to Methyl Eugenol by Male Bactrocera dorsalis (Diptera: Tephritidae)

Chieng AC, Hee AK, Wee SL

J Insect Sci, 2018 Sep 01;18(5).
PMID: 30351432 DOI: 10.1093/jisesa/iey104

The oriental fruit fly, Bactrocera dorsalis (Handel) is one of the most destructive pests of fruits. The discovery of methyl eugenol (ME) as a potent male attractant for this species has led to its successful use in area-wide fruit fly control programs such as male annihilation. While the antenna is recognized as primarily responsible for male flies' detection of attractants such as ME, little is known of the involvement of the maxillary palp. Using behavioral assays involving males with intact and ablated antennae and maxillary palp structures, we seek to ascertain the relative involvement of the maxillary palp in the ability of the male fly to detect ME. In cage bioassays (distance of ≤40 cm from the source), >97% of unmodified males will normally show a response to ME. Here, we showed that 17.6% of males with their antennae ablated were still attracted to ME versus 75.0% of males with their palps ablated. However, none of the antennae-ablated males were able to detect ME over a distance of >100 cm. Furthermore, wind tunnel bioassays showed that maxillary palp-ablated males took a significantly longer time compared to unablated males to successfully detect and eventually feed on ME. These results suggest that although the antennae are necessary for detection of ME over longer distances, at shorter distances, both antennae and maxillary palps are also involved in detecting ME. Hence, those palps may play a larger role than previously recognized in maneuvering males toward lure sources over shorter ranges.
Fulltext Historical perspective on the synonymization of the four major pest species belonging to the Bactrocera dorsalis species complex (Diptera, Tephritidae)

Hee AK, Wee SL, Nishida R, Ono H, Hendrichs J, Haymer DS, et al.

Zookeys, 2015.
PMID: 26798266 DOI: 10.3897/zookeys.540.6028

An FAO/IAEA-sponsored coordinated research project on integrative taxonomy, involving close to 50 researchers from at least 20 countries, culminated in a significant breakthrough in the recognition that four major pest species, Bactrocera dorsalis, Bactrocera philippinensis, Bactrocera papayae and Bactrocera invadens, belong to the same biological species, Bactrocera dorsalis. The successful conclusion of this initiative is expected to significantly facilitate global agricultural trade, primarily through the lifting of quarantine restrictions that have long affected many countries, especially those in regions such as Asia and Africa that have large potential for fresh fruit and vegetable commodity exports. This work stems from two taxonomic studies: a revision in 1994 that significantly increased the number of described species in the Bactrocera dorsalis species complex; and the description in 2005 of Bactrocera invadens, then newly incursive in Africa. While taxonomically valid species, many biologists considered that these were different names for one biological species. Many disagreements confounded attempts to develop a solution for resolving this taxonomic issue, before the FAO/IAEA project commenced. Crucial to understanding the success of that initiative is an accounting of the historical events and perspectives leading up to the international, multidisciplinary collaborative efforts that successfully achieved the final synonymization. This review highlights the 21 year journey taken to achieve this outcome.
Current knowledge of taxonomy, physiology and chemical ecology about Bactrocera dorsalis and its related species with comments to Wu et al. (2020)

Hee AK, Wee SL, Ono H, Nishida R, Tan KH

Mol Phylogenet Evol, 2021 03;156:107019.
PMID: 33248203 DOI: 10.1016/j.ympev.2020.107019
Fulltext Comparison of Knowledge, Attitude, and Practice among Communities Living in Hotspot and Non-Hotspot Areas of Dengue in Selangor, Malaysia

Ghani NA, Shohaimi S, Hee AK, Chee HY, Emmanuel O, Alaba Ajibola LS

Trop Med Infect Dis, 2019 Feb 15;4(1).
PMID: 30781369 DOI: 10.3390/tropicalmed4010037

BACKGROUND: Dengue has affected more than one-third of the world population and Malaysia has recorded an increase in the number of dengue cases since 2012. Selangor state recorded the highest number of dengue cases in Malaysia. Most of the dengue infections occur among people living in hotspot areas of dengue. This study aims to compare Knowledge, Attitude, and Practice among communities living in hotspot and non-hotspot dengue areas.
METHOD: Communities living in 20 hotspot and 20 non-hotspot areas in Selangor were chosen in this study where 406 participants were randomly selected to answer questionnaires distributed at their housing areas. Total marks of each categories were compared using t-test.
RESULT: Results show that there were significant mean differences in marks in Knowledge (p value: 0.003; 15.41 vs. 14.55) and Attitude (p value: < 0.001; 11.41 vs. 10.33), but not Practice (p value 0.101; 10.83 vs. 10.47) categories between communities of non-hotspot and hotspot areas. After considering two confounding variables which are education level and household income, different mean marks are found to be significant in Knowledge when education level acts as a covariate and Attitude when both act as covariates.
CONCLUSION: Overall results show that people living in non-hotspot areas had better knowledge and attitude than people living in hotspot areas, but no difference was found in practice. This suggests that public health education should be done more frequently with people with a low education background and low household income, especially in hotspot areas to fight dengue outbreak and make dengue cases decrease effectively.