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      North American Lauraceae: Terpenoid Emissions, Relative Attraction and Boring Preferences of Redbay Ambrosia Beetle, Xyleborus glabratus (Coleoptera: Curculionidae: Scolytinae)

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          Abstract

          The invasive redbay ambrosia beetle, Xyleborus glabratus, is the primary vector of Raffaelea lauricola, a symbiotic fungus and the etiologic agent of laurel wilt. This lethal disease has caused severe mortality of redbay ( Persea borbonia) and swampbay ( P. palustris) trees in the southeastern USA, threatens avocado ( P. americana) production in Florida, and has potential to impact additional New World species. To date, all North American hosts of X. glabratus and suscepts of laurel wilt are members of the family Lauraceae. This comparative study combined field tests and laboratory bioassays to evaluate attraction and boring preferences of female X. glabratus using freshly-cut bolts from nine species of Lauraceae: avocado (one cultivar of each botanical race), redbay, swampbay, silkbay ( Persea humilis), California bay laurel ( Umbellularia californica), sassafras ( Sassafras albidum), northern spicebush ( Lindera benzoin), camphor tree ( Cinnamomum camphora), and lancewood ( Nectandra coriacea). In addition, volatile collections and gas chromatography-mass spectroscopy (GC-MS) were conducted to quantify terpenoid emissions from test bolts, and electroantennography (EAG) was performed to measure olfactory responses of X. glabratus to terpenoids identified by GC-MS. Significant differences were observed among treatments in both field and laboratory tests. Silkbay and camphor tree attracted the highest numbers of the beetle in the field, and lancewood and spicebush the lowest, whereas boring activity was greatest on silkbay, bay laurel, swampbay, and redbay, and lowest on lancewood, spicebush, and camphor tree. The Guatemalan cultivar of avocado was more attractive than those of the other races, but boring response among the three was equivalent. The results suggest that camphor tree may contain a chemical deterrent to boring, and that different cues are associated with host location and host acceptance. Emissions of α-cubebene, α-copaene, α-humulene, and calamenene were positively correlated with attraction, and EAG analyses confirmed chemoreception of terpenoids by antennal receptors of X. glabratus.

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          Ethanol and (-)-alpha-Pinene: attractant kairomones for bark and ambrosia beetles in the southeastern US.

          In 2002-2004, we examined the flight responses of 49 species of native and exotic bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) to traps baited with ethanol and/or (-)-alpha-pinene in the southeastern US. Eight field trials were conducted in mature pine stands in Alabama, Florida, Georgia, North Carolina, and South Carolina. Funnel traps baited with ethanol lures (release rate, about 0.6 g/day at 25-28 degrees C) were attractive to ten species of ambrosia beetles (Ambrosiodmus tachygraphus, Anisandrus sayi, Dryoxylon onoharaensum, Monarthrum mali, Xyleborinus saxesenii, Xyleborus affinis, Xyleborus ferrugineus, Xylosandrus compactus, Xylosandrus crassiusculus, and Xylosandrus germanus) and two species of bark beetles (Cryptocarenus heveae and Hypothenemus sp.). Traps baited with (-)-alpha-pinene lures (release rate, 2-6 g/day at 25-28 degrees C) were attractive to five bark beetle species (Dendroctonus terebrans, Hylastes porculus, Hylastes salebrosus, Hylastes tenuis, and Ips grandicollis) and one platypodid ambrosia beetle species (Myoplatypus flavicornis). Ethanol enhanced responses of some species (Xyleborus pubescens, H. porculus, H. salebrosus, H. tenuis, and Pityophthorus cariniceps) to traps baited with (-)-alpha-pinene in some locations. (-)-alpha-Pinene interrupted the response of some ambrosia beetle species to traps baited with ethanol, but only the response of D. onoharaensum was interrupted consistently at most locations. Of 23 species of ambrosia beetles captured in our field trials, nine were exotic and accounted for 70-97% of total catches of ambrosia beetles. Our results provide support for the continued use of separate traps baited with ethanol alone and ethanol with (-)-alpha-pinene to detect and monitor common bark and ambrosia beetles from the southeastern region of the US.
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            The scent of a partner: ambrosia beetles are attracted to volatiles from their fungal symbionts.

            Invasive fungus-growing ambrosia beetles are an emerging threat to forest ecosystems and fruit industries, but management tools are lacking. Here we explored the potential of beetle symbionts-ambrosia fungi-as a source of attractants. Our focus was the redbay ambrosia beetle, Xyleborus glabratus, and its symbiotic fungus, Raffaelea lauricola, which are devastating lauraceous hosts in the southeastern United States. We also tested three additional co-occurring beetle species and their symbionts. Each beetle species was consistently attracted to the odors of its symbiotic fungal species, occasionally also to symbionts of other species, but never to non-symbiotic Trichoderma. We further confirmed attraction to ethanol (positive control) in some species. Thus, ambrosia fungi produce volatiles attractive to their vector beetles, which may have potential as novel lures for ambrosia beetle management.
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              What reaches the antenna? How to calibrate odor flux and ligand-receptor affinities.

              Physiological studies on olfaction frequently ignore the airborne quantities of stimuli reaching the sensory organ. We used a gas chromatography-calibrated photoionization detector to estimate quantities released from standard Pasteur pipette stimulus cartridges during repeated puffing of 27 compounds and verified how lack of quantification could obscure olfactory sensory neuron (OSN) affinities. Chemical structure of the stimulus, solvent, dose, storage condition, puff interval, and puff number all influenced airborne quantities. A model including boiling point and lipophilicity, but excluding vapor pressure, predicted airborne quantities from stimuli in paraffin oil on filter paper. We recorded OSN responses of Drosophila melanogaster, Ips typographus, and Culex quinquefasciatus, to known quantities of airborne stimuli. These demonstrate that inferred OSN tuning width, ligand affinity, and classification can be confounded and require stimulus quantification. Additionally, proper dose-response analysis shows that Drosophila AB3A OSNs are not promiscuous, but highly specific for ethyl hexanoate, with other earlier proposed ligands 10- to 10 000-fold less potent. Finally, we reanalyzed published Drosophila OSN data (DoOR) and demonstrate substantial shifts in affinities after compensation for quantity and puff number. We conclude that consistent experimental protocols are necessary for correct OSN classification and present some simple rules that make calibration, even retroactively, readily possible.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                9 July 2014
                : 9
                : 7
                : e102086
                Affiliations
                [1 ]United States Department of Agriculture, Agricultural Research Service, Subtropical Horticulture Research Station, Miami, Florida, United States of America
                [2 ]Archbold Biological Station, Lake Placid, Florida, United States of America
                [3 ]United States Department of Agriculture, Forest Service, Southern Research Station, Asheville, North Carolina, United States of America
                [4 ]United States Department of Agriculture, Forest Service, Forest Health Protection, Stanislaus National Forest, Sonora, California, United States of America
                [5 ]United States Department of Agriculture, Agricultural Research Service, Beltsville Area Research Center, Electron and Confocal Microscopy Unit, Beltsville, Maryland, United States of America
                [6 ]University of Florida, Tropical Research and Education Center, Homestead, Florida, United States of America
                Swedish University of Agricultural Sciences, Sweden
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: PK WM JN. Performed the experiments: PK WM JN GB. Analyzed the data: PK JN NE. Contributed reagents/materials/analysis tools: PK JN GP AM MM MD GB RP NE. Contributed to the writing of the manuscript: PK.

                Article
                PONE-D-14-16327
                10.1371/journal.pone.0102086
                4090202
                25007073
                a17f7731-4b98-4320-9dc0-95a1eb691d30
                Copyright @ 2014

                This is an open-access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 14 April 2014
                : 15 June 2014
                Page count
                Pages: 13
                Funding
                Funding was provided by the USDA-ARS National Plant Disease Recovery System and the Florida Avocado Administrative Committee. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Agriculture
                Crops
                Crop Diseases
                Fruits
                Pests
                Insect Pests
                Weevils
                Forestry
                Ecology
                Behavioral Ecology
                Chemical Ecology
                Custom metadata
                The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.

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