Olfactory responses of Drosophila suzukii females to host plant volatiles

Drosophila suzukii Matsumura, an endemic pest in southeast Asia, has invaded Europe and the U.S.A. Unlike most of its closely related sibling species, the serrated ovipositor of D. suzukii permits ovipositing in undamaged fresh fruits. In the present study, volatiles are identified from host plants that are potentially involved in D. suzukii host recognition and oviposition behaviour. It is shown that mated females are attracted to volatiles emitted from intact fruits. The antennally‐active suite of compounds released from the fresh fruits is identified by gas chromatography coupled with electroantennographic detection, as well as gas chromatography‐mass spectrometry. In olfactometer bioassays, mated females are significantly attracted to an electroantennographically active volatile, isoamyl acetate, when tested at 10 µg of synthetic compound in a rubber septa, which has a release rate comparable to that of fresh fruits. In addition, a genomic survey shows that D. suzukii not only possesses the full repertoire of genes encoding odorant receptors activated by isoamyl acetate in D. melanogaster, but also that one of the genes, OR67a, is represented by five duplicated copies. These results indicate that D. suzukii uses olfactory cues to select oviposition sites. The identification of volatiles emitted by host fruits that attract D. suzukii may aid in the development of a selective and efficient synthetic lure for monitoring this pest. As a close relative of Drosophila melanogaster, D. suzukii provides a unique opportunity for understanding the physiological mechanisms involved in the shift of this species from use of rotten to ripe fruits for oviposition.     

水果害虫斑翅果蝇体内微生物菌群研究进展

斑翅果蝇是一种在全球范围内造成危害的重要水果害虫,其主要分布于亚、美、欧三大洲。斑翅果蝇的产卵器可以刺破水果表皮,将卵产在未完全成熟的水果中,卵孵化为幼虫后,幼虫取食水果,直接降低产量,从而对水果产业造成损失。近年以来,越来越多的研究表明昆虫微生物对宿主昆虫影响很大。例如昆虫微生物可以调控寄主昆虫的生长发育、个体适应性及生殖等。昆虫与其共生微生物间的关系成为昆虫生物学研究的热点内容。本文综述了近些年关于斑翅果蝇微生物多样性的研究,探讨了微生物菌群及内共生菌Wolbachia对斑翅果蝇生长发育、行为、生殖、抗病毒等的影响,以便为寻找控制斑翅果蝇种群的策略提供参考依据。     

Chemical cues from competitors change the oviposition preference of Drosophila suzukii

Although Drosophila suzukii Matsumura (Diptera: Drosophilidae) infests fresh and ripening fruits, it is attracted to fermented fruits as well. Because fermented fruits attract other flies too, if D. suzukii utilizes fermented fruits as oviposition substrates, competition can be more intense on them. To avoid such competition, D. suzukii may change oviposition preference when particular species of competitor flies are present, but the effect of odor cues associated with competitors on the oviposition preference of D. suzukii is still unknown. To examine such an effect, we investigated the oviposition preference of D. suzukii in the presence of four competitor fly species – Drosophila melanogaster Meigen, Drosophila lutescens Malloch, Drosophila rufa Kikkawa & Peng, and Drosophila auraria Peng – and D. suzukii itself. We prepared artificial substrates with yeast treatment (Y⁺: yeast supplementation, Y⁻: control) and competitor fly treatment (F⁺: pre-inoculated with competitor fly odor, F⁻: control), and performed two-choice experiments using the substrates with various Y and F treatments. Our results showed that D. suzukii oviposited more eggs on Y⁺ substrates than on Y⁻ substrates when no competitor flies were present and the presence of competitor flies influenced D. suzukii’s oviposition preference for yeast-supplemented substrates and its effect changed depending on the competitor fly species. If the presence of competitors around fallen fruits on the ground suppresses D. suzukii’s oviposition on the fallen fruits and facilitates the oviposition on non-fermenting substrates in nature, it may drive D. suzukii to use ripening fruits on the tree. Such selective pressure may facilitate the evolution of morphological traits such as a serrated ovipositor in D. suzukii.     

Identification of active components from volatiles of Chinese bayberry, <Emphasis Type="Italic">Myrica rubra</Emphasis> attractive to <Emphasis Type="Italic">Drosophila suzukii</Emphasis>

Drosophila suzukii (Diptera: Drosophilidae) is native to Southeast Asia and now has become a severe pest of several soft fruits in Europe and the Americas. It causes considerable damage to Chinese bayberry, Myrica rubra, in China. In the present study, we employed gas chromatograph–electroantennographic detection (GC–EAD) together with behavioural bioassays and trapping experiments to identify volatile semiochemicals emitted by Chinese bayberry attracting D. suzukii. Electrophysiological experiments revealed the presence of six EAD-active compounds from ripe bayberry fruits, including methyl (E)-3-hexenoate, methyl (E)-2-hexenoate, ethyl (E)-2-hexenoate, α-ylangene, α-humulene and an unidentified compound that elicited consistent antennal response. In two-choice bioassays, bayberry fruits attracted all responding flies, and significantly more flies responded to the volatile extract of bayberry fruits. Four EAD-active compounds were attractive to mated female D. suzukii at lower doses (0.01 and 0.1 µg), but showed repellency at higher doses (10 and 100 µg). Mixtures of these four compounds at different ratios attracted D. suzukii flies at all test doses (0.1, 1 and 10 µg). Both male and female flies were trapped by a mixture of synthetic methyl (E)-3-hexenoate, methyl (E)-2-hexenoate, ethyl (E)-2-hexenoate and α-humulene in a ratio of 1:1.3:1:6.4 in the field trapping experiment. Significantly more males than females were captured in the trap baited with the synthetic blend, and the percentages of D. suzukii captured out of all flies by the traps baited with lure were higher than that baited with blank control. Our findings may provide insights into the olfactory responses of D. suzukii to specific host plant volatiles, and contribute to further development of an effective lure for monitoring D. suzukii in the field.     

Innate Olfactory Responses of <Emphasis Type="Italic">Asobara japonica</Emphasis> Toward Fruits Infested by the Invasive Spotted Wing Drosophila

Insect parasitoids are often manipulated to improve biological control programs for various arthropod pests. Volatile compounds can be a relevant cue used by most parasitoid hymenoptera for host or host microhabitat location. Here, we studied olfactory responses of the braconid Asobara japonica Belokobylskij, an Asiatic endoparasitoid of the invasive pest Drosophila suzukii (Matsumura), toward its host and host substrates. Adult A. japonica displayed an innate attraction to undescribed volatile cues from infested host fruits irrespectively of the juvenile rearing experience, i.e. they respond to a novel cue subsequently used for microhabitat selection. These data suggest that A. japonica parasitoids mass-reared on artificial diet and factitious host (D. melanogaster) can successfully locate their hosts. Naïve female parasitoids did not show a preference towards any of the tested host media. However, the enforced adult experience with the rearing host medium modified the olfactory preference patterns toward non-natal host fruits. These findings provide evidence of associative learning during the adult stage of A. japonica, and demonstrate its plasticity in exploiting the volatiles from various fruits infested by D. suzukii.     

A chitinase from Euphorbia characias latex is a novel and powerful plant‐based pesticide against Drosophila suzukii

Drosophila suzukii attacks on developing soft fruits have recently caused important economic losses in Europe. This study explores the effectiveness of a new control strategy against this insect pest that is based on a plant chitinase extracted from the latex of the Mediterranean spurge, Euphorbia characias. The ability of the purified Euphorbia latex chitinase (ELC) to degrade the chitin exoskeleton of D. suzukii was assessed using confocal laser scanning microscopy. ELC treatment caused reduced larval growth, higher mortality and notable degradation of external insect structures. Therefore, the chitinase may induce a double effect on the D. suzukii larvae, a direct injury on the larval bodies and an action as antifeedant. The effects of the ELC treatment were also tested on leaves of the insect's host plants, Fragaria × ananassa and Rubus idaeus, using physiological parameters (chlorophyll concentration, chlorophyll fluorescence, leaf gas exchange and water potential) and defence gene expression (FaPGIP, FaChi2_1 and FaChi2_2) as stress indicators. ELC at concentrations effective against D. suzukii did not damage the host plants. Only plant defence gene expression was somewhat enhanced during the early hours after ELC application. In conclusion, ELC, a natural product, proved to be an effective tool for use in the development of an environmentally friendly integrated management strategy against D. suzukii, a pest whose control by conventional chemical insecticides is problematic.     

RESPONSE OF PARASITOID MICROPLITIS MEDIATOR TO PLANT VOLATILES IN AN OLFACTOMETER*

Abstract The behavioral responses of Microplitis mediator were measured in a four‐armed olfactometer. Leaves extract of the suitable host plant of its insect host, cotton, elicited higher olfactory responses; while leaves extract of the less suitable host plant of its insect host, tomato or tobacco, elicited moderate responses. Volatiles from the least suitable host plant of its insect host, hot pepper, elicited the lowest responses. The different preferences to four plant leaves extracts suggested that M. mediator could discriminate between suitable and less suitable host plants of its insect host by semiochemicals. Green leaf volatiles (GLVs), cis‐3‐hexen‐1‐yl acetate, hexanal, and nonan‐1‐ol, are the most attractive compound among the 7 tested compounds. The parasitoids preferred host‐damaged plant to undamaged plant, suggesting that they were able to distinguish between host and non‐host of their insect host through varying chemical blends. Experience with cotton leaves extract prior to experiment enhanced their responses to the same odor.     

A New Synthetic Method of Methyl Ketones

A simple and stereospecific method has been developed for the synthesis of cis-jasmone. 3-Methyl-2-cyclopentenone (V) was condensed with cis-2-pentenyl chloride, which was prepared from propargyl alcohol, by means of sodium amide in liquid ammonia to yield stereochemically pure cis-jasmone (VIb) in 39~40% yield. By the similar direct alkylation, trans-jasmone (VIc) and dihydrojasmone (VIa) were also synthesized in moderate yields.     

Odor discrimination using insect electroantennogram responses from an insect antennal array

Insects have a highly developed olfactory sensory system, mainly based in their antennae, for the detection and discrimination of volatile compounds in the environment. Electroantennogram (EAG) response profiles of five different insect species, Drosophila melanogaster, Heliothis virescens, Helicoverpa zea, Ostrinia nubilalis and Microplitis croceipes, showed different, species-specific EAG response spectra to 20 volatile compounds tested. The EAG response profiles were then reconstructed for each compound across the five insect species. Most of the compounds could be distinguished by comparing the response spectra. We then used a four-antenna array, called a Quadro-probe EAG, to see if we could discriminate among odorants based on the relative EAG amplitudes evoked when the probe was placed in plumes in a wind tunnel and in a field. Stable EAG responses could be simultaneously and independently recorded with four different insect antennae mounted on the Quadro-probe, and different volatile compounds could be distinguished in real time by comparing relative EAG responses with a combination of differently tuned insect antennae. Regardless of insect species or EAG amplitudes, antennae on the Quadro-probe maintained their responsiveness with higher than 1 peak/s of time resolution.     

Seasonal monitoring of Drosophila suzukii and its non-crop hosts in Wuhu,Eastern China

Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) is a serious pest that prefers fresh fruits and is native to Southeast Asia. In our study, apple cider vinegar bait traps were used to capture and monitor the population dynamics of this native pest in Wuhu City, China, from May/June 2017 to May 2018. The research was conducted at 15 locations in two fruit orchards in Wuhu. Traps caught more adults in general in a Meiling blueberry orchard than in a Xicun mixed orchard, and the highest trap counts occurred near harvest time (October). Females had more mature eggs from September to November, and the number of mature eggs declined thereafter. We found several non-crop hosts, which can provide food and reproductive resources for D. suzukii and are common in forests and field margins. By comparing the number of captured adults in the Meiling and Xicun orchards, we found that blueberry was preferred by D. suzukii among the fruits in our search. Fruit ripening times differed among crops; therefore, fly populations moved between crop and non-crop habitats during the year or had varying population dynamics on different crops in different seasons. The D. suzukii population and the number of mature eggs decreased in summer and winter but increased in spring and autumn. Drosophila suzukii had higher survival rates with blueberry than those with other fruits, and D. suzukii could use four non-crop species growing around the orchards as host plants.     

<Emphasis Type="Italic">Drosophila suzukii</Emphasis> in Southern Neotropical Region: Current Status and Future Perspectives

Non-native insect pests are often responsible for important damage to native and agricultural plant hosts. Since Drosophila suzukii Matsumura (Diptera: Drosophilidae) has become an important pest in North America and Europe (i.e., in 2008), the global production of soft thin-skinned fruits has faced severe production losses. In the southern Neotropical region, however, the first record of D. suzukii occurred in 2013 in the south of Brazil. It has also been recorded in Uruguay, Argentina, and Chile. Despite its recent occurrence in the southern Neotropical region, the fast dispersion of D. suzukii has inspired local research efforts in an attempt to mitigate the consequences of this insect pest invasion. In this forum, we explore the current status of D. suzukii in southern Neotropical regions, discussing its future perspectives. Additionally, we attempt to draft activities and a research agenda that may help to mitigate the losses caused by D. suzukii in native and commercial soft-skinned fruits produced in this region. Currently, D. suzukii appears to be well established in the south of Brazil, but considering the entire southern Neotropical region, the invasion panorama is still underinvestigated. The lack of studies and regulatory actions against D. suzukii has contributed to the invasion success of this species in this region. Considering several peculiarities of both the pest biology and the environmental of this region, the authors advocate for the need of intensive and integrative studies toward the development and implementation of area-wide integrated pest management programs against D. suzukii in the southern Neotropical region.     

Behavioral evidence for contextual olfactory‐mediated avoidance of the ubiquitous phytopathogen Botrytis cinerea by Drosophila suzukii

Herbivorous insects may benefit from avoiding the smell produced by phytopathogens infecting plant host tissue if the infected tissue reduces insect fitness. However, in many cases the same species of phytopathogen can also infect host plant tissues that do not directly affect herbivore fitness. Thus, insects may benefit from differentiating between pathogen odors emanating from food and nonfood tissues. This is based on the hypothesis that unnecessarily staying attentive to pathogen odor from nonfood tissue may incur opportunity costs associated with not responding to other important survival functions. In this study adults of Drosophila suzukii Matsumura, an invasive larval frugivore, showed reduced attraction to the odor of raspberry fruit, a food tissue, when infected with Botrytis cinerea Pers., a ubiquitous phytopathogen, in favor of odors of uninfected raspberry fruit. Moreover, D. suzukii oviposited fewer eggs on infected raspberry fruit relative to uninfected raspberry fruit. Larval survival and adult size after eclosion were significantly reduced when reared on B. cinerea‐infected raspberry relative to uninfected fruit. Interestingly, when the behavioral choice experiment was repeated using Botrytis‐infected vs. ‐uninfected strawberry leaves, a nonfood tissue, in combination with fresh raspberry fruit, odor from B. cinerea‐infected leaves did not reduce D. suzukii attraction to raspberries relative to raspberries with uninfected leaves. These behavioral results illustrate the important role context can play in odor‐mediated interactions between insects, plants and microbes. We discuss implications of our findings for developing a repellent that can be useful for the management of D. suzukii.     

Improving monitoring tools for spotted wing drosophila,Drosophila suzukii

Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) were trapped in the field using colored plastic sphere traps coated with insect Tangle‐trap. Red and black spheres captured significantly more D. suzukii than white spheres. Translucent deli‐cup traps deployed in cherry orchards and baited with yeast, the Alpha Scents lure, or the Scentry lure captured significantly more flies than the Trécé lure and Suzukii bait; all attractants had poor selectivity for D. suzukii. No‐choice evaluations of attractants conducted in field cages corroborated the cherry orchard field study, though translucent deli‐cup traps provisioned with the yeast bait captured significantly more flies than those baited with the Alpha Scents lure. Red sphere traps baited with the Scentry lure captured 3–6× more flies than the deli‐cup trap baited with the same lure, and 3–4× more flies than the deli‐cup trap baited with yeast bait, demonstrating that a trap integrating both visual and olfactory cues is a superior tool for monitoring D. suzukii. Moreover, this simple sticky, dry trap design requires far less labor and maintenance than does a liquid‐based deli‐cup trap.     

Electroantennogram responses of a parasitic wasp, Microplitis croceipes, to host-related volatile and anthropogenic compounds

The parasitic wasp Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) showed its own characteristic electroantennogram (EAG) response profiles to 13 host‐related (cis‐3‐hexenol, α‐pinene (R)‐(+)‐limonene (S)‐(–)‐limonene, trans‐β‐ocimene (±)‐linalool, (–)‐trans‐caryophyllene, α‐humulene, nerolidol, trans‐nerolidol, cis‐nerolidol, methyl jasmonate and indole) and four anthropogenic (2‐diisopropylaminoethanol, 2,2′‐thiodiethanol, 2‐methyl‐5‐nitroaniline and cyclohexanone) volatile compounds. These profiles were similar between males and females except for 2‐diisopropylaminoethanol, which elicited significantly larger EAG responses in males. Among the compounds tested, cis‐3‐hexenol, linalool and cyclohexanone elicited the largest EAG responses. EAG responses were not influenced by the age of wasps between 1 and 13 days after emergence. EAG responses were dose‐dependent, and highly EAG‐active compounds elicited significant EAG responses with less than 10 μg of the compounds at source. Quantification of compounds released from an odour cartridge indicates that release rate is highly dependent on the chemical nature of stimuli, showing up to 10 000‐fold differences in the amount released between different compounds when the same amount was loaded in the odour cartridge. Wasps having undergone a behavioural training regime to be attracted to either cyclohexanone or methyl jasmonate did not show any differences in EAG responses from those of untrained wasps.     

Nutritional geometry and fitness consequences in Drosophila suzukii,the Spotted‐Wing Drosophila

Since its arrival to North America less than a decade ago, the invasive Spotted‐Wing Drosophila (Drosophila suzukii) has inflicted substantial economic losses on soft fruit agriculture due to its ability to oviposit into ripening fruits. More effective management approaches for this species are needed, but little is known about the factors that influence behavioral choices made by D. suzukii when selecting hosts, or the consequences that their offspring experience when developing in different environments. Using a nutritional geometry methodology, we found that the ratio of proteins‐to‐carbohydrates (P:C) present in media greatly influenced adult D. suzukii behavior and subsequent offspring development. Whereas adult flies showed a strong bias in their oviposition and association behaviors toward carbohydrate‐rich foods, larval survival and eclosion rate were strongly dependent on protein availability. Here, we explore the preference–performance hypothesis (PPH), in which females are predicted to oviposit on medias that provide the greatest offspring benefits, in regard to its relevance in D. suzukii behavior and consequences for management. Our results provide valuable insight into the ecology and evolution of this species that may hopefully lead to more effective management strategies.     

Potential host fruits for Drosophila suzukii: olfactory and oviposition preferences and suitability for development

Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), the spotted wing drosophila, is a pest endemic to Southeast Asia that invaded the Americas and Europe in 2008. In contrast to most of its congeners, D. suzukii possesses a serrated ovipositor that allows it to lay eggs in unwounded commercial fruits, resulting in severe revenue losses for the industry. The purpose of this study was to determine the susceptibility of known host fruits, including cherry, strawberry, blueberry, and grape, and potential host fruits, such as banana and apple, to attack by D. suzukii. Based on the responses to volatile cues offered in a six‐choice olfactometer, the preference of female D. suzukii was ranked in the following order: strawberry = cherry > banana = apple = blueberry = grape, but in no‐choice and choice oviposition tests, the preferences were ranked as follows: cherry > strawberry = blueberry > grape = banana > apple. Furthermore, we reconfirmed that D. suzukii mainly targets rotten fruit for feeding and ripe fruit for oviposition, and females preferred fruits with intensive mechanical damage. Based on developmental parameters, apple was the least suitable host. This study has implications for the control of D. suzukii, especially in mixed fruit orchards, by providing a promising avenue for exploiting behaviour‐based control tools and emphasizing the importance of phenology in host fruit susceptibility.     

Morphological and ultrastructural characterization of olfactory sensilla in Drosophila suzukii: Scanning and transmission electron microscopy

Drosophila suzukii is a serious horticultural and quarantine pest, damaging various berry crops. Although the active use of olfactory communication in D. suzukii is well-known, their olfactory sensory system has not been comprehensively reported. Therefore, the present study was carried out to understand the morphology, distribution and ultrastructure of olfactory sensilla present in the antennae and maxillary palps of D. suzukii, through scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The olfactory sensilla on the antennae of D. suzukii in both sexes could be classified into three major morphological types, basiconic, trichoid and coeloconic sensilla, according to their shapes. The antennal basiconic sensilla were further divided into three subtypes and the antennal trichoid sensilla into two subtypes, respectively, according to the size of individual sensillum. In contrast to the antennal olfactory sensilla showing diverse morphology, basiconic sensilla was the only type of olfactory sensilla in the maxillary palps of D. suzukii. The basiconic sensilla in the maxillary palps could be further classified into three subtypes, based on their size. Our SEM and TEM observations indicated that multiple nanoscale pores are present on the surface of all types of olfactory sensilla in the antennae and maxillary palps, except coeloconic sensilla. The difference in the morphological types and the distribution of olfactory sensilla suggests that their olfactory functions are different between antennae and maxillary palps in D. suzukii. The results of this study provide useful information for further studies to determine the function of olfactory sensilla in D. suzukii and to understand their chemical communication system.     

Invasive host for invasive pest: when the Asiatic cherry fly (Drosophila suzukii) meets the American black cherry (Prunus serotina) in Europe

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Screening of 25 different natural crop protection products against Drosophila suzukii

Drosophila suzukii is a major pest of soft‐skinned fruits, and insecticides are often used to prevent fruit damage caused by oviposition. As D. suzukii produces many generations per year, repeated insecticide applications are required. Furthermore, D. suzukii attacks ripening and ripe fruits shortly before harvest. Therefore, the use of synthetic insecticides is limited by long pre‐harvest intervals and maximum residue limits. To be able to offer producers immediate and sustainable solutions, we tested 25 natural crop protection products with three different application methods in a laboratory screening. We show that application method is an important factor for the efficacy of the tested products. Of six natural insecticides, only Spinosad was toxic for D. suzukii and reduced the oviposition on treated blueberries. The tested oil products had no control effect and products based on different entomopathogenic fungi and Bacillus thuringiensis rather enhanced oviposition. Mineral products (Kaolin, CaCO₃, Ca(OH)₂ and clinoptilolith) applied as spray solutions were not toxic, but significantly reduced oviposition on blueberries. We provide the first study in which different application methods have been used to compare numerous, commercially available, natural crop protection products with different modes of action against adult D. suzukii. Our findings provide consultants and producers with important insights for the development of sustainable pest control strategies against D. suzukii.     

The influence of monoterpenoids and phenol derivatives on Drosophila melanogaster viability

The present study investigated the effect of mono/bicyclic terpenoids and phenol derivatives on the viability of Drosophila melanogaster and their influence on the multiplication of the nuclear genome. The fertility and viability of fruit flies were assessed after oral administration and inhalation exposure of compounds 1–5: guaiacol, eugenol, borneol, menthol and carvacrol. The influence of terpenoids and phenols on the degree of chromosomes polyteny in salivary gland cells of D. melanogaster larvae was determined. Among all tested compounds, carvacrol demonstrated the most significant impact on fecundity and insect survival when inhaled or adding to the culture medium. Oral administration of carvacrol had an impact on giant chromosomes increasing their average level of chromosome polyteny degree while eugenol adding to culture medium had the opposite effect. The possible mechanism of terpenoids and phenols action is discussed.