Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores

Host plant selection by a monophagous herbivore is not mediated by quantitative changes in unique plant chemistry : Agonopterix alstroemeriana and Conium maculatum

Host plant selection by ovipositing females is a key process determining the success of phytophagous insects. In oligophagous lepidopterans, host-specific plant secondary chemicals are expected to be dominant factors governing oviposition behavior; distinctive compounds can serve as high-contrast signals that clearly differentiate confamilial hosts from non-hosts increasing the accuracy of host quality evaluation. Agonopterix alstroemeriana (Clerk) (Lepidoptera: Oecophoridae) and Conium maculatum L. (Apiaceae) form an extremely specialized plant-herbivore system, with A. alstroemeriana monophagous on C. maculatum, a plant with few other insect herbivores at least in part due to its virtually unique capacity among plants to produce piperidine alkaloids. Here we have studied the response of A. alstroemeriana oviposition to unique host plant secondary metabolites, piperidine alkaloids, and widespread compounds, mono- and sesquiterpenes, in a concentration-dependent fashion. Rates of oviposition were negatively correlated with Z-ocimene concentrations. To confirm the deterrent properties of this monoterpene for A. alstroemeriana oviposition, we conducted a choice experiment using artificially damaged C. maculatum plants, with higher emission of volatiles, and undamaged control plants. Damaged plants were less preferred as oviposition sites compared to the controls. The lack of association between oviposition and piperidine alkaloids, defenses unique to Conium species, suggests that quantitative changes of these species-specific chemicals do not play a predominant role in host selection by the monophagous A. alstroemeriana

Geographical variation in alkaloid production in Conium maculatum populations experiencing differential herbivory by Agonopterix alstroemeriana

Conium maculatum, a Eurasian weed naturalized in North America, contains high concentrations of piperidine alkaloids, which act as chemical defenses against herbivores. In the United States, C. maculatum was largely free from herbivory until approximately 30 years ago, when it was re-associated via accidental introduction with a monophagous European herbivore, the oecophorid caterpillar Agonopterix alstroemeriana. At present, A. alstroemeriana is found in a continuum of re-association time and intensities with C. maculatum across the continent; in the Pacific Northwest, A. alstroemeriana can cause severe damage, resulting in some cases in complete defoliation. Studies in biological control and invasion biology have yet to determine whether plants re-associated with a significant herbivore from the area of indigeneity increase their chemical defense investment in areas of introduction. In this study, we compared three locations in the U.S. (New York, Washington and Illinois) where C. maculatum experiences different levels of herbivory by A. alstroemeriana to determine the association between the intensity of the interaction, as measured by damage, and chemical defense production. Total alkaloid production in C. maculatum was positively correlated with A. alstroemeriana herbivory levels; plants from New York and Washington, with higher herbivory levels, invested two and four times more N to alkaloid synthesis than did plants from Illinois. Individual plants with lower concentrations of alkaloids from a single location in Illinois experienced more damage by A. alstroemeriana, suggestive of a preference on the part of the insect for plants with less chemical defense. These results suggest that A. alstroemeriana may act either as a selective agent or inducing agent for C. maculatum and increase its toxicity in its introduced range

Genetic variation of alkaloid production in Conium maculatum after reassociation with the specialist moth Agonopterix alstroemeriana

Conium maculatum, a Eurasian weed naturalized in North America, contains high concentrations of piperidine alkaloids. In the United States, C. maculatum was largely free from herbivory until approximately 30 years ago, when it was re-associated via accidental introduction with a monophagous European herbivore, the oecophorid caterpillar Agonopterix alstroemeriana. At present, A. alstroemeriana is found in a continuum of re-association time and intensities with C. maculatum across the continent; in the Pacific Northwest, A. alstroemeriana can cause severe damage, resulting in some cases in complete defoliation. Studies in biological control and invasion biology have yet to determine whether plants re-associated with a significant herbivore from the area of indigeneity increase their chemical defense investment in areas of introduction. In this study we compared the secondary chemistry of C. maculatum in three locations in the U.S. (New York, Washington and Illinois) where C. maculatum experiences different levels of herbivory by A. alstroemeriana. We conducted an in situ sampling and a common garden experiment to determine the association between the intensity of the interaction, as measured by damage, and chemical defense production. Total alkaloid production in C. maculatum was positively correlated with A. alstroemeriana herbivory levels. Plants from NY and WA, with higher herbivory levels, invested two and four times more N to alkaloid synthesis, respectively, than did plants from IL. Individual plants with higher concentrations of monoterpenes and alkaloids received fewer eggs and experienced less damage by A. alstroemeriana suggestive of a preference on the part of the insect for plants with lower levels of chemical defense. These results suggest that A. alstroemeriana may act as a selective agent for C. maculatum and prolonged reassociation may increase toxicity of this noxious weed in its introduced range

A reference case for economic evaluations in osteoarthritis: An expert consensus article from the European Society for Clinical and Economic Aspects of Osteoporosis and Osteoarthritis (ESCEO)

Background: General recommendations for a reference case for economic studies in rheumatic diseases were published in 2002 in an initiative to improve the comparability of cost-effectiveness studies in the field. Since then, economic evaluations in osteoarthritis (OA) continue to show considerable heterogeneity in methodological approach. Objectives: To develop a reference case specific for economic studies in OA, including the standard optimal care, with which to judge new pharmacologic and non-pharmacologic interventions. Methods: Four subgroups of an ESCEO expert working group on economic assessments (13 experts representing diverse aspects of clinical research and/or economic evaluations) were charged with producing lists of recommendations that would potentially improve the comparability of economic analyses in OA: outcome measures, comparators, costs and methodology. These proposals were discussed and refined during a face-to-face meeting in 2013. They are presented here in the format of the recommendations of the recently published Consolidated Health Economic Evaluation Reporting Standards (CHEERS) statement, so that an initiative on economic analysis methodology might be consolidated with an initiative on reporting standards. Results: Overall, three distinct reference cases are proposed, one for each hand, knee and hip OA; with diagnostic variations in the first two, giving rise to different treatment options: interphalangeal or thumb-based disease for hand OA and the presence or absence of joint malalignment for knee OA. A set of management strategies is proposed, which should be further evaluated to help establish a consensus on the "standard optimal care" in each proposed reference case. The recommendations on outcome measures, cost itemisation and methodological approaches are also provided. Conclusions: The ESCEO group proposes a set of disease-specific recommendations on the conduct and reporting of economic evaluations in OA that could help the standardisation and comparability of studies that evaluate therapeutic strategies of OA in terms of costs and effectiveness

Larval case architecture and implications of host-plant associations for North American Coleophora (Lepidoptera; Coleophoridae)

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The comparative responsiveness of Hospital Universitario Princesa Index and other composite indices for assessing rheumatoid arthritis activity

Objective To evaluate the responsiveness in terms of correlation of the Hospital Universitario La Princesa Index (HUPI) comparatively to the traditional composite indices used to assess disease activity in rheumatoid arthritis (RA), and to compare the performance of HUPI-based response criteria with that of the EULAR response criteria. Methods Secondary data analysis from the following studies: ACT-RAY (clinical trial), PROAR (early RA cohort) and EMECAR (pre-biologic era long term RA cohort). Responsiveness was evaluated by: 1) comparing change from baseline (Delta) of HUPI with Delta in other scores by calculating correlation coefficients; 2) calculating standardised effect sizes. The accuracy of response by HUPI and by EULAR criteria was analyzed using linear regressions in which the dependent variable was change in global assessment by physician (Delta GDA-Phy). Results Delta HUPI correlation with change in all other indices ranged from 0.387 to 0.791); HUPI's standardized effect size was larger than those from the other indices in each database used. In ACT-RAY, depending on visit, between 65 and 80% of patients were equally classified by HUPI and EULAR response criteria. However, HUPI criteria were slightly more stringent, with higher percentage of patients classified as non-responder, especially at early visits. HUPI response criteria showed a slightly higher accuracy than EULAR response criteria when using Delta GDA-Phy as gold standard. Conclusion HUPI shows good responsiveness in terms of correlation in each studied scenario (clinical trial, early RA cohort, and established RA cohort). Response criteria by HUPI seem more stringent than EULAR''s

Host plant selection by a monophagous herbivore is not mediated by quantitative changes in unique plant chemistry : Agonopterix alstroemeriana and Conium maculatum

Host plant selection by ovipositing females is a key process determining the success of phytophagous insects. In oligophagous lepidopterans, host-specific plant secondary chemicals are expected to be dominant factors governing oviposition behavior; distinctive compounds can serve as high-contrast signals that clearly differentiate confamilial hosts from non-hosts increasing the accuracy of host quality evaluation. Agonopterix alstroemeriana (Clerk) (Lepidoptera: Oecophoridae) and Conium maculatum L. (Apiaceae) form an extremely specialized plant-herbivore system, with A. alstroemeriana monophagous on C. maculatum, a plant with few other insect herbivores at least in part due to its virtually unique capacity among plants to produce piperidine alkaloids. Here we have studied the response of A. alstroemeriana oviposition to unique host plant secondary metabolites, piperidine alkaloids, and widespread compounds, mono- and sesquiterpenes, in a concentration-dependent fashion. Rates of oviposition were negatively correlated with Z-ocimene concentrations. To confirm the deterrent properties of this monoterpene for A. alstroemeriana oviposition, we conducted a choice experiment using artificially damaged C. maculatum plants, with higher emission of volatiles, and undamaged control plants. Damaged plants were less preferred as oviposition sites compared to the controls. The lack of association between oviposition and piperidine alkaloids, defenses unique to Conium species, suggests that quantitative changes of these species-specific chemicals do not play a predominant role in host selection by the monophagous A. alstroemeriana