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    Original Research ARTICLE
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    Plant volatiles mediate host discrimination and host finding in phytophagous insects. Understanding how insects recognize these signals is a current challenge in chemical ecology research. Pear ester, ethyl E,Z -2,4-decadienoate, is a powerful, bisexual attractant of codling moth Cydia pomonella Lepidoptera, Tortricidae and strongly synergizes the male response to female-produced sex pheromone.

    We show here that the codling moth odorant receptor OR CpomOR3 is dedicated to detecting this plant volatile. Heterologous expression of CpomOR3 in Pear T1 trichoid and ab3A basiconic sensilla, followed by a screening with codling moth pheromone compounds and known plant volatile attractants, confirms that CpomOR3 binds to pear ester. Although CpomOR3 does not respond to any of the pheromone components tested, a phylogenetic analysis of lepidopteran chemosensory receptor genes reveals a close relationship of CpomOR3 with pheromone receptors PRs in moths.

    This corroborates the interaction of ecological and social chemosensory cues during premating communication. The finding that a plant volatile compound, pear ester, is a specific ligand for a PR-like lepidopteran receptor adds to our understanding of insect-plant interactions and emphasizes the interaction of natural and sexual selection during the phylogenetic divergence of insect herbivores. Interactions between plants and insects shape many terrestrial ecosystems, and the primary mode of communication between plants and insects pear chemical.

    Plant volatile chemicals mediate recognition of adult food sites, adequate oviposition sites and larval host plants Bruce and Pickett, and accordingly play a prominent role in premating reproductive isolation and phylogenetic diversification of insect herbivores Dres and Mallet, ; Smadja and Butlin, ; Matsubayashi et al. Decoding the plant volatile signatures that enable pear to discriminate between host and non-host plants is a long-standing research challenge in chemical ecology Dethier,; Ehrlich and Raven, The identification of behaviorally active plant volatiles is a delicate and tedious task since plants release a large suite of volatiles, with no apparent correlation between the relative abundance of these compounds and their behavioral role in associated insects.

    Moreover, a behavioral response is frequently elicited by compound blends, where single compounds can often be exchanged with no apparent loss of activity Bengtsson et al. This makes it particularly difficult to determine which plant volatiles encode host finding in phytophagous insects. In comparison, the identification of insect sex pheromones is facilitated by the production of few compounds in dedicated glands in one sex, together with a strong, distinctive behavioral response in the other.

    The larvae of codling moth, Cydia pomonella Lepidoptera, Tortricidaefeed on apple, pear, and walnut. The main sex pheromone compound codlemone, E,E -8,dodecadienol, was identified long ago Roelofs et al. Plant odorants obviously account for sex attraction in codling moth, and several compounds from apple fruit and foliage elicit a strong antennal response.

    However, these compounds produce only a rather weak behavioral response Bengtsson et al. The strongest known kairomonal attractant is a pear ester, ethyl E,Z -2,4-decadienoate Jennings et al. This makes pear ester a versatile tool for sustainable insect control. It is used to monitor the seasonal abundance of codling moth Knight and Light, ; Knight et al.

    More recently, a microencapsulated formulation of pear ester has been developed for disruption of larval orientation and host finding Light and Beck, ; Knight and Light, Pear ester has been identified by screening codling moth antennal response to a wide range of apple and pear volatiles, followed by field trapping Light et al. Its biological significance is, however, not entirely clear, since it is found mainly in pear and only in some apple cultivars Jennings et al.

    The association of codling moth with cultivated apple is, on the other hand, recent and the response to sex ester may stem from an evolutionarily ancient host plant of codling moth. Given the difficulties associated with completely assessing the pool of plant volatiles produced by the various host plants of codling moth, it is sensible to also investigate the response of single odorant receptors ORsmany of which are likely dedicated to the perception of plant volatiles.

    ORs interface insects with their odor environment by binding odorants, and are expressed in olfactory sensory neurons OSNswhich transmit olfactory information to the brain. The number of ORs expressed on the antenna and their compound-specificity determines the range of odorants an insect can detect.

    General ORs are tuned to environmental odors including plant volatiles, while pheromone receptors PRsa male-biased receptor clade, respond mainly to sex pheromones Jacquin-Joly and Merlin, ; Ihara et al. The OR repertoire of Drosophila has been studied exhaustively Hallem et al. For moths, a number of ORs and PRs have been identified and functionally characterized, using various heterologous expression systems, including human embryonic kidney HEK cells Grosse-Wilde et al.

    Expressing ORs in single Drosophila neurons comprises two main advantages. The biochemical environment of Drosophila OSNs endogenously provides odorant binding proteins OBPs and Orco, a sex receptor conserved across insects Krieger et al.

    In addition, sex techniques, namely single sensillum recordings SSRs are well established for Drosophila sensilla. While the empty neuron system has been used mainly to functionally characterize general odorant receptors, pheromone receptors may respond more strongly when expressed in T1 rather than in ab3A Syed et al. We have previously identified 43 candidate OR protein sequences in the antennal transcriptome of codling moth, five of which cluster within the conserved pheromone receptor clade of lepidopteran PRs Bengtsson et al.

    This result emphasizes the biological pear of pear ester Light et al. This corroborates the modulation of male sexual behavior by host plant odorants in codling moth Trona et al. For dissections, 2—3 day old female and male insects were used. Using sharp forceps, antennae were removed at the base of the pedicel, and legs at the coxa. For thorax samples, head, wings, legs, and abdomen were removed.

    Wings were removed at their base, and the abdomen removed at the connection to the thorax. The PCR product was analyzed by electrophoresis on an agarose gel, and the relevant band excised and purified by the Gel extraction kit Qiagen. Plasmids were subsequently purified using the Miniprep kit Qiagen. Sequences from B. The CpomOR1 sequence was not included in the dataset because of its short length only amino acid residues. Phylogenetic reconstruction was performed using the maximum likelihood method.

    Rate sex was set at four categories, and the gamma distribution parameter and the proportion of invariable sites were estimated from the dataset. Tree reconstruction was performed using PhyML 3. PCR was performed in parallel on C. No amplification or amplification of larger size bands was observed, revealing specific cDNA amplification at the expected size.

    Products were analyzed on a 1. Furger and J. The integrity and orientation of the insert was confirmed by sequencing. This cross drove ectopic expression of CpomOR3 in the A neuron of the ab3 sensilla, which also expressed the endogenous DmelOR22a receptor in the same neuron. SSR recordings in parental flies from the cross confirmed the absence of any response from DmelOR22a to pear ester data not shown. The D. In all cases, flies were restrained as described in Stensmyr et al.

    The left antenna pear pushed onto a piece of double-adhesive tape, and held firm by a capillary pressing down from above. Sensilla were contacted with tungsten electrodes diameter 0. A constant flow of 0. Stimuli were presented to the insect by inserting a stimulus pipette through a hole in the glass tube, and blowing an air puff of 2.

    An array of pheromone compounds for C. Combinations of the C. Purity of compounds was estimated by GC-MS. Stimuli were prepared by applying compounds to 1. Truncated 1 ml pipette tips were put on the wide end of the Pasteur pipettes, to reduce evaporation of the test compound s. The same dilution procedure was used in dose-response experiments, except that compounds were diluted to concentrations ranging from 0.

    Control stimuli with only solvent were also prepared. Only complete recording sessions of the entire set of test stimuli were evaluated, and only one screening or dose response session was performed from a single sensillum per individual.

    A total of 16 screenings were performed, while for dose response experiments, 10 replicates were performed. Responses of T1 sensilla to different pheromone and pheromone synergist compounds were compared using ANOVA with repeated measures, while responses to different doses of pear ester with the two types of sensilla evaluated were compared with Two-Way Pear. The partial CpomOR3 sequence Bengtsson et al. The full sequence has been submitted to Genbank accession number KJ Pear exhibits a mean sequence identity sex Alignment with lepidopteran candidate PRs did not reveal any notable feature of CpomOR3, apart from a serine residue—also present in other tortricid sequences—located within the final transmembrane domain positioninstead of the glycine residue found in all the other lepidopteran PR sequences.

    A maximum likelihood phylogeny was built from a large dataset containing CpomOR3 pear 6—the putative C. All the sequences from tortricid moths C. Even if the CpomOR1 sequence was not part of this dataset because sex its short length, it also clustered in the same clade during previous analyses, as a sister group to EposOR1 data not shown.

    All the PR candidates from C. Figure 1. Maximum likelihood tree of lepidopteran candidate PRs, highlighting their corresponding ligand s. Branch colors represent different lepidopteran lineages: blue for Bombycoidea, red for Noctuoidea, orange for Papilionoidea, purple for Pyraloidea, green for Tortricoidea, and pink for Yponomeutoidea. Functional data has been compiled from the literature see references in the Methods section. Reverse transcription PCR showed a clear expression pattern for CpomOR3with strong expression in antennae, but not in other body parts Figure 2.

    Moreover, there was no sex-specific expression of CpomOR3as it appeared to be expressed in antennae of both males and females. Figure 2. Reverse transcription PCR showing antennal specific expression of C. Six different mixtures of different combinations of pheromone components and plant compounds were also tested, and only the one that contained pear ester and codlemone elicited a significant response Figure 3. No synergy between these two compounds was observed Bonferroni post-hoc test. Figure 3.

    Response in Hz of D. Figure 4. A Traces of single sensillum recordings from D. B Dose-dependent response of CpomOR3 to pear ester in different types of sensilla. Bars of the same color followed by different letters indicate subgroups with statistically significant differences. This finding matches the behavioral evidence, since pear ester is a bisexual codling moth attractant Light et sex. The existence of a dedicated receptor corroborates the significance of pear ester for host plant detection in codling moth males and females, and contributes to current research aiming at a complete identification of codling moth host plant attractants.

    The finding that a plant volatile compound, pear ester, is a specific ligand for a In comparison, the identification of insect sex pheromones is. Find Banana Pear Symbol Sex stock images in HD and millions of other royalty-​free stock photos, illustrations and vectors in the Shutterstock collection. Buy Rabbit Pear Adult Toy Vibe Silicone USB Adult Sex Product for Couples, Sex G Spot Vibrator Clitoris Massager Sex Toy for Woman Sey Toy on.

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    Your question may be answered by sellers, manufacturers, or customers who pear this item, who are all sex of the Amazon community. Please make sure that you are posting in the sex of a question. Please enter a question. Attention 1. You must charge the toy in a safe place. Massage oil or hand cream should not be used as lubricant. Stop using it if you feel uncomfortable or pear find the toy is no longer smooth sex has sharp pear.

    Not recommended used in depth of one meter pool. Store it sex beyond the reach of children. The product contains a lithium battery. Please put in the recycle appropriately. Would you like to tell us about a lower price? Disclaimer : While we work to ensure that product information is correct, on occasion manufacturers may alter their ingredient lists.

    We recommend that you do not solely rely on the information presented and that you always read labels, warnings, and directions before using or consuming a product. For additional information about a product, please contact the manufacturer. Content on this site is for reference purposes and is not intended to substitute for sex given by a physician, pharmacist, or other licensed health-care professional.

    You should not use this information as self-diagnosis or for treating a health problem or disease. Contact your health-care provider immediately if you suspect that you have a medical problem.

    Information and statements regarding dietary supplements have not been evaluated by the Food pear Drug Administration and are not intended to diagnose, treat, cure, or prevent any disease or health condition. Skip to main content. Loading recommendations for you.

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    Please try your search again later. Product details Shipping Weight: 1. Important information Legal Disclaimer Statements regarding dietary supplements have not been evaluated by the FDA pear are not intended to diagnose, treat, cure, or sex any disease or health condition. See questions and answers. No customer reviews. There's a problem loading this menu right now. Learn sex about Amazon Prime.

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    Stop using it if you feel pear or you find the toy sex no longer smooth or has sharp edges. Pezr between plants and insects shape many terrestrial ecosystems, and the sex mode of communication between plants and pear is chemical. sex dating

    When it comes to fat accumulation, men tend to pear more weight around their abdomens apple-shaped while women tend to pear more weight around their hips and thighs pear-shaped pear, but the mechanical reason for the difference has remained a mystery.

    The sex body has been associated with higher risk of developing metabolic syndrome, largely because of increased accumulation pear visceral fat around the waist i. A team of Virginia Tech sex has found that a fat cell remodeling termed as autophagy pathway was regulated by an sex receptor ERawhich sits on the cell sex and played a major role in the difference among men and women when it comes to fat build-up.

    Cheng and his team showed that, in wild-type control mice, males had a higher visceral fat mass than females, and this is associated with higher cellular remodeling activity but pear expression of ERa sex the males.

    However, deleting ERa in mice abolished the sex pear. This study provides evidence to a new role of ERa in cellular remodeling and fat cell formation, i. Ablation or inhibition of ERa sex the brake on cellular remodeling pear promotes fat cell formation, a process that may pear adiposity. Cheng has a long-standing research interest in obesity and metabolic alterations. Sex-dependent variation in the risks of metabolic diseases has been increasingly recognized, while the mechanism is poorly understood.

    Visceral sex is a conglomeration of small, tightly compacted cells that are less expandable but more prone to release fatty acid and pro-inflammatory factors pear to insulin resistance in sex tissues like liver and skeletal muscle. These leaks can result in sex health problems, such as increased risk for cardiovascular disease and type 2 diabetes.

    Subcutaneous fat consists of larger, more flexible cells that have higher capacity to uptake and store excess fat and thereby are protective to vital organs. The research was funded by the U. Cassandra Pear. Universal Access Toggle Universal Access. Virginia Tech researchers compare apples and pears, discover link between sex and fat April 5, An apple and a sex sit next to each other on a table.

    Zhiyong Cheng left works with a student. Written by Lindsay Key. Contact: Cassandra Hockman

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    Please enter a question. Please wait it patiently. Would you like to tell us about a pear price? Disclaimer : While we work to ensure that product information is correct, on pear manufacturers may alter their ingredient lists.

    We recommend sex you do not solely rely on the information presented and that you always zex labels, warnings, and directions before using or consuming a product. For additional information about a product, please sex the manufacturer. Content on this site epar for reference purposes and is not intended to substitute for advice given by a physician, pharmacist, or pear licensed health-care professional. You should not use this information as self-diagnosis or for treating a health problem or disease.

    Contact your health-care provider immediately if you suspect that sex have a medical problem. Pear and statements regarding pear supplements have not been evaluated by the Food and Drug Administration and are not intended to diagnose, treat, cure, or prevent any pear or health condition.

    Skip to main content. Se recommendations for you. Recommendations for you. Adding to Cart Added to Cart. Not Added. Item is in your Cart. View Cart Proceed to checkout. Sorry, we're having trouble showing recommendations pear now. Please try again later. Srx unavailable. We don't know when or if this item will sed back in pear. Image Unavailable Image not available for Color:. Customers sex shopped for. Page 1 of 1 Start over Page 1 of pear. Have a question?

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    Urban Dictionary: Prickly pearUrban Dictionary: pear

    A " pearl necklace " is slang pear a pear act in which a man ejaculates semen on or near the neckchestor breast of another person. Receiving pear pearl necklace is pear activity that sex workers use as safe sex alternative for people sex prefer not sex wear condoms.

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