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Environmental exposure to cold is increasingly being associated with changes in metabolism. We developed and tested the hypothesis that exposure to cold drives systemic effects in lipid metabolism. Specifically, (i) that energy storage and provision adapts to the cold by altering triglyceride distribution and (ii) that membranes adapt to cold conditions by becoming more unsaturated. These hypotheses were designed to identify the underlying mechanisms that govern the response of mammalian systems to cold. To test these hypotheses, we used a metabolic network analysis. An established model of cold exposure was used, from which lipidomics data that represents the system was collected. The network analysis showed that triglyceride metabolism is altered on exposure to cold, with several smaller effects that are not straightforward, such as changes to the abundance and distribution of odd chain fatty acids. The range and profile of phosphatidylcholine and phosphatidylinositol were modified, but there was little change in phosphatidylethanolamine or sphingomyelin. These results support the hypothesis, and show that exposure to cold is a system-wide phenomenon that requires or drives changes across a range of metabolic pathways., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 James et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Lipid metabolism is recognised as being central to growth, disease and health. Lipids, therefore, have an important place in current research on globally significant topics such as food security and biodiversity loss. However, answering questions in these important fields of research requires not only identification and measurement of lipids in a wider variety of sample types than ever before, but also hypothesis-driven analysis of the resulting 'big data'. We present a novel pipeline that can collect data from a wide range of biological sample types, taking 1 000 000 lipid measurements per 384 well plate, and analyse the data systemically. We provide evidence of the power of the tool through proof-of-principle studies using edible fish (mackerel, bream, seabass) and colonies of Bombus terrestris . Bee colonies were found to be more like mini-ecosystems and there was evidence for considerable changes in lipid metabolism in bees through key developmental stages. This is the first report of either high throughput LCMS lipidomics or systemic analysis in individuals, colonies and ecosystems. This novel approach provides new opportunities to analyse metabolic systems at different scales at a level of detail not previously feasible, to answer research questions about societally important topics.

Lablab ( Lablab purpureus L.) is an important food and livestock feed legume that can also enhance soil fertility. However, its production is limited by insect pests, notably the black bean aphid ( Aphis fabae ). The present field study was conducted to determine the difference in the contribution of lablab genotypes and natural field margin vegetation (FMV) to the abundance and diversity of natural enemies and the damage, incidence, and abundance of bean aphids. Eighteen lablab genotypes were planted in the presence or absence of FMV in a randomized complete block design experiment replicated four times. Data on aphid abundance, incidence, and severity of damage were collected at four growth stages of the crop. Lablab genotypes significantly influenced aphid incidence, suggesting some level of tolerance to aphid colonization. Findings showed that lablab genotypes were a significant influence on natural enemy species richness with no statistical difference for abundance and natural enemy species diversity. However, the genotypes did not vary significantly in their influence on the number of aphid natural enemies. FMV was associated with low bean aphid damage. Overall, the presence or absence of FMV did not influence the number of natural enemies caught on the crop. This concurs with recent work that shows a similar number of natural enemies with field margin plants but may reflect the reduced number of pest insects. Cropping seasons influenced aphid abundance and damage severity, with the populations developing at the early stages of lablab development and decreasing as the crop advanced. This pattern was similar both in the presence or absence of FMV. The findings of this study highlight the important contribution of crop genotype together with the presence of field margin species in the regulation of aphids and their natural enemies in lablab., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision., (Copyright © 2024 Karimi, Nyaanga, Mulwa, Ogendo, Bett, Cheruiyot, Arnold, Belmain and Stevenson.)

Main Conclusion: One of seven Solanum taxa studied displayed associations between pollen presence and floral scent composition and volume, suggesting buzz-pollinated plants rarely use scent as an honest cue for foraging pollinators. Floral scent influences the recruitment, learning, and behaviour of floral visitors. Variation in floral scent can provide information on the amount of reward available or whether a flower has been visited recently and may be particularly important in species with visually concealed rewards. In many buzz-pollinated flowers, tubular anthers opening via small apical pores (poricidal anthers) visually conceal pollen and appear similar regardless of pollen quantity within the anther. We investigated whether pollen removal changes floral scent composition and emission rate in seven taxa of buzz-pollinated Solanum (Solanaceae). We found that pollen removal reduced both the overall emission of floral scent and the emission of specific compounds (linalool and farnesol) in S. lumholtzianum. Our findings suggest that in six out of seven buzz-pollinated taxa studied here, floral scent could not be used as a signal by visitors as it does not contain information on pollen availability., (© 2024. Crown.)

Fall armyworm, Spodoptera frugiperda (FAW) is a cosmopolitan crop pest species that has recently become established in sub-Saharan Africa and Southeast Asia. Current FAW control is almost entirely dependent on synthetic pesticides. Biopesticides offer a more sustainable alternative but have limitations. For example, pyrethrum is an effective botanical insecticide with low mammalian toxicity but is highly UV labile, resulting in a rapid loss of efficacy in the field. Beauveria bassiana is an entomopathogenic fungus that is more persistent, but there is a time lag of several days before it causes insect mortality and leads to effective control. The combination of these biopesticides could mitigate their drawbacks for FAW control. Here we evaluated the efficacy of pyrethrum and B. bassiana as individual treatments and in combination against 3 rd instar FAW. Four different combinations of these two biopesticides were tested, resulting in an antagonistic relationship at the lowest concentrations of B. bassiana and pyrethrum (1 × 10 4 conidia mL -1 with 25 ppm) and an additive effect for the other 3 combined treatments (1 × 10 4 conidia mL -1 with 100 ppm and 1 × 10 5 conidia mL -1 with 25 ppm and 100 ppm pyrethrum). Additionally, a delay in efficacy from B. bassiana was observed when combined with pyrethrum as well as a general inhibition of growth on agar plates. These results appear to show that this particular combination of biopesticides is not universally beneficial or detrimental to pest control strategies and is dependent on the doses of each biopesticide applied. However, the additive effect shown here at specific concentrations does indicate that combining biopesticides could help overcome the challenges of persistence seen in botanical pesticides and the slow establishment of EPF, with the potential to improve effectiveness of biopesticides for IPM., (© 2024. The Author(s).)

Specialization in plant pollination systems can arise from traits that function as filters of flower visitors. This may involve chemical traits such as floral volatiles that selectively attract favoured visitors and non-volatile nectar constituents that selectively deter disfavoured visitors through taste or longer-term toxic effects or both. We explored the functions of floral chemical traits in the African milkweed Gomphocarpus physocarpus, which is pollinated almost exclusively by vespid wasps, despite having nectar that is highly accessible to other insects such as honeybees. We demonstrated that the nectar of wasp-pollinated G. physocarpus contains cardenolides that had greater toxic effects on Apis mellifera honeybees than on Vespula germanica wasps, and also reduced feeding rates by honeybees. Behavioural experiments using natural compositions of nectar compounds showed that these interactions are mediated by non-volatile nectar chemistry. We also identified volatile compounds with acetic acid as a main component in the floral scent of G. physocarpus that elicited electrophysiological responses in wasp antennae. Mixtures of these compounds were behaviourally effective for attraction of V. germanica wasps. The results show the importance of both volatile and non-volatile chemical traits as filters that lead to specialization in plant pollination systems., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Little is known concerning terpenoids produced by members of the fungal order Ophiostomales, with the member Harringtonia lauricola having the unique lifestyle of being a beetle symbiont but potentially devastating tree pathogen. Nine known terpenoids, including six labdane diterpenoids ( 1 - 6 ) and three hopane triterpenes ( 7 - 9 ), were isolated from H. lauricola ethyl acetate (EtOAc) extracts for the first time. All compounds were tested for various in vitro bioactivities. Six compounds, 2 , 4 , 5 , 6 , 7 , and 9 , are described functionally. Compounds 2 , 4 , 5 , and 9 expressed potent antiproliferative activity against the MCF-7, HepG2 and A549 cancer cell lines, with half-maximal inhibitory concentrations (IC 50 s) ~12.54-26.06 μM. Antimicrobial activity bioassays revealed that compounds 4 , 5 , and 9 exhibited substantial effects against Gram-negative bacteria ( Escherichia coli and Ralstonia solanacearum ) with minimum inhibitory concentration (MIC) values between 3.13 and 12.50 μg/mL. Little activity was seen towards Gram-positive bacteria for any of the compounds, whereas compounds 2 , 4 , 7 , and 9 expressed antifungal activities ( Fusarium oxysporum ) with MIC values ranging from 6.25 to 25.00 μg/mL. Compounds 4 , 5 , and 9 also displayed free radical scavenging abilities towards 2,2-diphenyl-1-picrylhydrazyl (DPPH) and superoxide (O 2- ), with IC 50 values of compounds 2 , 4 , and 6 ~3.45-14.04 μg/mL and 22.87-53.31 μg/mL towards DPPH and O 2- , respectively. These data provide an insight into the biopharmaceutical potential of terpenoids from this group of fungal insect symbionts and plant pathogens.

Leaf traits often vary with plant neighbourhood composition, which in turn may mediate plant susceptibility to herbivory. However, it is unknown whether there are any common patterns of change in leaf trait expression in response to neighbourhood diversity, and whether these responses confer increased resistance or susceptibility to herbivores.We used meta-analysis to combine data from 43 studies that examined the influence of neighbourhood diversity on eight physical and chemical leaf traits that could affect herbivory. All leaf traits apart from leaf thickness were highly plastic and exhibited significant differences between plant monocultures and species mixtures, but the direction of effect was variable. Leaf toughness was the only trait that displayed a significant decrease with plant diversity, whereas specific leaf area (SLA) and leaf nitrogen were both marginally increased in species mixtures.The magnitude and direction of leaf trait responses to neighbourhood diversity were independent of plant density and phylogenetic diversity, but changes in SLA correlated positively with plant species richness. SLA was also significantly increased in experimental studies, but not in observational studies, while neighbourhoods containing nitrogen-fixers were associated with increased leaf nitrogen and reduced phenolics. When studies on the over-represented species Betula pendula were removed from the analysis, the effect of neighbourhood diversity on leaf toughness became nonsignificant, but phenolics were significantly reduced in diverse neighbourhoods composed of mature trees, and marginally reduced in species mixtures across all studies.Increases in plant neighbourhood diversity are often associated with reductions of herbivory, although in some cases, the reverse occurs, and plants growing in species mixtures are found to suffer greater herbivory than those in monocultures. This study offers a potential explanation for the latter phenomenon, as our results show that leaf trait expression is highly plastic in response to neighbourhood diversity, and in certain cases could lead to increased leaf quality, which in turn could promote greater rates of herbivory. Read the free Plain Language Summary for this article on the Journal blog., Competing Interests: Julia Koricheva is an associate editor of Functional Ecology, but took no part in the peer review and decision‐making processes for this paper., (© 2023 The Authors. Functional Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

The insect pathogenic fungus, Ascosphaera apis , is the causative agent of honeybee chalk brood disease. Amylases are secreted by many plant pathogenic fungi to access host nutrients through the metabolism of starch, and the identification of new amylases can have important biotechnological applications. Production of amylase by A. apis in submerged culture was optimized using the response surface method (RSM). Media composition was modeled using Box-Behnken design (BBD) at three levels of three variables, and the model was experimentally validated to predict amylase activity ( R 2 = 0.9528). Amylase activity was highest (45.28 ± 1.16 U/mL, mean ± SE) in media composed of 46 g/L maltose and1.51 g/L CaCl 2 at a pH of 6.6, where total activity was ~11-fold greater as compared to standard basal media. The enzyme was purified to homogeneity with a 2.5% yield and 14-fold purification. The purified enzyme had a molecular weight of 75 kDa and was thermostable and active in a broad pH range (> 80% activity at a pH range of 7-10), with optimal activity at 55 °C and pH = 7.5. Kinetic analyses revealed a K m of 6.22 mmol/L and a V max of 4.21 μmol/mL·min using soluble starch as the substrate. Activity was significantly stimulated by Fe 2+ and completely inhibited by Cu 2+ , Mn 2+ , and Ba 2+ (10 mM). Ethanol and chloroform (10% v / v ) also caused significant levels of inhibition. The purified amylase essentially exhibited activity only on hydrolyzed soluble starch, producing mainly glucose and maltose, indicating that it is an endo-amylase (α-amylase). Amylase activity peaked at 99.38 U/mL fermented in a 3.7 L-bioreactor (2.15-fold greater than what was observed in flask cultures). These data provide a strategy for optimizing the production of enzymes from fungi and provide insight into the α-amylase of A. apis .

Sterols have several roles in planta, including as membrane components. Sterols are also essential nutrients for insects. Based on this, and the different functions of leaves and pollen, we tested the hypotheses that (a) the sterolome is different in leaves and pollen from the same plant, (b) pollens from wind- and insect pollinated plants comprise different sterols, and (c) sterol provision in pollen-rewarding angiosperms differs from nectar-rewarding species. A novel approach to sterolomics was developed, using LCMS to determine the sterol profile of leaf and pollen from a taxonomically diverse range of 36 plant species. Twenty-one sterols were identified unambiguously, with several more identified in trace amounts. C 29 sterols dominated the sterolome in most plants. The sterol composition was significantly different in leaf and pollen and their main sterols evolved in different ways. The sterolome of pollen from animal- and wind-pollinated was also significantly different, but not between nectar- and pollen-rewarding species. Our results suggest that the sterol composition in different plant tissues is linked to their biological functions. Sterol composition in pollen might be driven by physical role rather than the nutrient needs of pollinating insects., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Background: Bees provide essential pollination services for many food crops and are critical in supporting wild plant diversity. However, the dietary landscape of pollen food sources for social and solitary bees has changed because of agricultural intensification and habitat loss. For this reason, understanding the basic nutrient metabolism and meeting the nutritional needs of bees is becoming an urgent requirement for agriculture and conservation. We know that pollen is the principal source of dietary fat and sterols for pollinators, but a precise understanding of what the essential nutrients are and how much is needed is not yet clear. Sterols are key for producing the hormones that control development and may be present in cell membranes, where fatty-acid-containing species are important structural and signalling molecules (phospholipids) or to supply, store and distribute energy (glycerides)., Aim of the Review: In this critical review, we examine the current general understanding of sterol and lipid metabolism of social and solitary bees from a variety of literature sources and discuss implications for bee health., Key Scientific Concepts of Review: We found that while eusocial bees are resilient to some dietary variation in sterol supply the scope for this is limited. The evidence of both de novo lipogenesis and a dietary need for particular fatty acids (FAs) shows that FA metabolism in insects is analogous to mammals but with distinct features. Bees rely on their dietary intake for essential sterols and lipids in a way that is dependent upon pollen availability., (© 2023. The Author(s).)

Grayanotoxin I (GTX I) is a major toxin in leaves of Rhododendron species, where it provides a defence against insect and vertebrate herbivores. Surprisingly, it is also present in R. ponticum nectar, and this can hold important implications for plant-pollinator mutualisms. However, knowledge of GTX I distributions across the genus Rhododendron and in different plant materials is currently limited, despite the important ecological function of this toxin. Here we characterise GTX I expression in the leaves, petals, and nectar of seven Rhododendron species. Our results indicated interspecific variation in GTX I concentration across all species. GTX I concentrations were consistently higher in leaves compared to petals and nectar. Our findings provide preliminary evidence for phenotypic correlation between GTX I concentrations in defensive tissues (leaves and petals) and floral rewards (nectar), suggesting that Rhododendron species may commonly experience functional trade-offs between herbivore defence and pollinator attraction., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Botanical insecticides offer an environmentally benign insect pest management option for field crops with reduced impacts on natural enemies of pests and pollinators while botanically rich field margins can augment their abundance. Here, we evaluated the non-target effects on natural enemies and pest control efficacy on bean aphids in Lablab of three neem- and pyrethrum-based botanical insecticides (Pyerin75EC®, Nimbecidine® and Pyeneem 20EC®) and determine the influence of florally rich field margin vegetation on the recovery of beneficial insects after treatment. The botanical insecticides were applied at the early and late vegetative growth stages. Data were collected on aphids (abundance, damage severity and percent incidence) and natural enemy (abundance) both at pre-spraying and post-spraying alongside Lablab bean yield. The efficacy of botanical insecticides was similar to a synthetic pesticide control and reduced aphid abundance by 88% compared with the untreated control. However, the number of natural enemies was 34% higher in botanical insecticide-treated plots than in plots treated with synthetic insecticide indicating that plant-based treatments were less harmful to beneficial insects. The presence of field margin vegetation increased further the number of parasitic wasps and tachinid flies by 16% and 20%, respectively. This indicated that non-crop habitats can enhance recovery in beneficial insect populations and that botanical insecticides integrate effectively with conservation biological control strategies. Higher grain yields of 2.55-3.04 and 2.95-3.23 t/ha were recorded for both botanical insecticide and synthetic insecticide in the presence of florally enhanced field margins in consecutive cropping seasons. Overall, these data demonstrated that commercial botanical insecticides together with florally rich field margins offer an integrated, environmentally benign and sustainable alternative to synthetic insecticides for insect pest management and increased productivity of the orphan crop legume, Lablab., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (© 2022 The Authors. Journal of Applied Entomology published by Wiley‐VCH GmbH.)

Field margins support important ecosystem services including natural pest regulation. We investigated the influence of field margins on the spatial and temporal distribution of natural enemies (NEs) of bean pests in smallholder farming systems. We sampled NEs from high and low plant diversity bean fields using sweep netting and coloured sticky traps, comparing monocropped and intercropped farms. NEs collected from within crops included predatory bugs, lacewings, predatory flies, parasitic flies, parasitic wasps, lady beetles, and a range of other predatory beetles; with the most dominant group being parasitic wasps. Overall, high plant diversity fields had a higher number of NEs than low-diversity fields, regardless of sampling methods. The field margin had a significantly higher number of lacewings, parasitic wasps, predatory bugs, syrphid flies, and other predatory beetles relative to the crop, but beneficial insects were collected throughout the fields. However, we observed marginally higher populations of NEs in intercropping than in monocropping although the effect was not significant in both low and high plant diversity fields. We recommend smallholder farmers protect the field margins for the added benefit of natural pest regulation in their fields.

Evidence from the last few decades indicates that pollinator abundance and diversity are at risk, with many species in decline. Anthropogenic impacts have been the focus of much recent work on the causes of these declines. However, natural processes, from plant chemistry, nutrition and microbial associations to landscape and habitat change, can also profoundly influence pollinator health. Here, we argue that these natural processes require greater attention and may even provide solutions to the deteriorating outlook for pollinators. Existing studies also focus on the decline of individuals and colonies and only occasionally at population levels. In the light of this we redefine pollinator health and argue that a top-down approach is required focusing at the ecological level of communities. We use examples from the primary research, opinion and review articles published in this special issue to illustrate how natural processes influence pollinator health, from community to individuals, and highlight where some of these processes could mitigate the challenges of anthropogenic and natural drivers of change. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

Plant compounds associated with herbivore defence occur widely in floral nectar and can impact pollinator health. We showed previously that Rhododendron ponticum nectar contains grayanotoxin I (GTX I) at concentrations that are lethal or sublethal to honeybees and a solitary bee in the plant's non-native range in Ireland. Here we further examined this conflict and tested the hypotheses that nectar GTX I is subject to negative pollinator-mediated selection in the non-native range, but that phenotypic linkage between GTX I levels in nectar and leaves acts as a constraint on independent evolution. We found that nectar GTX I experienced negative directional selection in the non-native range, in contrast to the native Iberian range, and that the magnitude and frequency of pollinator limitation indicated that selection was pollinator-mediated. Surprisingly, nectar GTX I levels were decoupled from those of leaves in the non-native range, which may have assisted post-invasion evolution of nectar without compromising the anti-herbivore function of GTX I (here demonstrated in bioassays with an ecologically relevant herbivore). Our study emphasizes the centrality of pollinator health as a concept linked to the invasion process, and how post-invasion evolution can be targeted toward minimizing lethal or sub-lethal effects on pollinators. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

Antimicrobial nectar secondary metabolites can support pollinator health by preventing or reducing parasite infections. To better understand the outcome of nectar metabolite-parasite interactions in pollinators, we determined whether the antiparasitic activity was altered through chemical modification by the host or resident microbiome during gut passage. We investigated this interaction with linden ( Tilia spp.) and strawberry tree ( Arbutus unedo ) nectar compounds . Unedone from A. unedo nectar inhibited the common bumblebee gut parasite Crithidia bombi in vitro and in Bombus terrestris gynes. A compound in Tilia nectar, 1-[4-(1-hydroxy-1-methylethyl)-1,3-cyclohexadiene-1-carboxylate]-6- O -β-d-glucopyranosyl-β-d-glucopyranose (tiliaside), showed no inhibition in vitro at naturally occurring concentrations but reduced C. bombi infections of B. terrestris workers. Independent of microbiome status, tiliaside was deglycosylated during gut passage, thereby increasing its antiparasitic activity in the hindgut, the site of C. bombi infections . Conversely, unedone was first glycosylated in the midgut without influence of the microbiome to unedone-8- O -β-d-glucoside, rendering it inactive against C. bombi , but subsequently deglycosylated by the microbiome in the hindgut, restoring its activity. We therefore show that conversion of nectar metabolites by either the host or the microbiome modulates antiparasitic activity of nectar metabolites. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

Objective: Polyunsaturated fatty acid (PUFA) supplements have been trialled as a treatment for a number of conditions and produced a variety of results. This variety is ascribed to the supplements, that often comprise a mixture of fatty acids, and to different effects in different organs. In this study, we tested the hypothesis that the supplementation of individual PUFAs has system-level effects that are dependent on the molecular structure of the PUFA., Methods: We undertook a network analysis using Lipid Traffic Analysis to identify both local and system-level changes in lipid metabolism using publicly available lipidomics data from a mouse model of supplementation with FA(20:4n-6), FA(20:5n-3), and FA(22:6n-3); arachidonic acid, eicosapentaenoic acid, and docosahexaenoic acid, respectively. Lipid Traffic Analysis is a new computational/bioinformatics tool that uses the spatial distribution of lipids to pinpoint changes or differences in control of metabolism, thereby suggesting mechanistic reasons for differences in observed lipid metabolism., Results: There was strong evidence for changes to lipid metabolism driven by and dependent on the structure of the supplemented PUFA. Phosphatidylcholine and triglycerides showed a change in the variety more than the total number of variables, whereas phosphatidylethanolamine and phosphatidylinositol showed considerable change in both which variables and the number of them, in a highly PUFA-dependent manner. There was also evidence for changes to the endogenous biosynthesis of fatty acids and to both the elongation and desaturation of fatty acids., Conclusions: These results show that the full biological impact of PUFA supplementation is far wider than any single-organ effect and implies that supplementation and dosing with PUFAs require a system-level assessment., (Copyright © 2022 The Author(s). Published by Elsevier GmbH.. All rights reserved.)

Wild bee populations are declining due to human activities, such as land use change, which strongly affect the composition and diversity of available plants and food sources. The chemical composition of food (i.e., nutrition) in turn determines the health, resilience, and fitness of bees. For pollinators, however, the term 'health' is recent and is subject to debate, as is the interaction between nutrition and wild bee health. We define bee health as a multidimensional concept in a novel integrative framework linking bee biological traits (physiology, stoichiometry, and disease) and environmental factors (floral diversity and nutritional landscapes). Linking information on tolerated nutritional niches and health in different bee species will allow us to better predict their distribution and responses to environmental change, and thus support wild pollinator conservation., Competing Interests: Declaration of interests The authors declare no conflicts of interest., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Flower-rich field margins provide habitats and food resources for natural enemies of pests (NEs), but their potential, particularly in the tropics and on smallholder farms, is poorly understood. We surveyed field margins for plant-NE interactions in bean fields. NEs most often interacted with Bidens pilosa (15.4% of all interactions) and Euphorbia heterophylla (11.3% of all interactions). In cage trials with an aphid-infested bean plant and a single flowering margin plant, the survival of Aphidius colemani, the most abundant parasitoid NE in bean fields, was greater in the presence of Euphorbia heterophylla than Bidens pilosa , Tagetes minuta , and Hyptis suaveolens . UV-fluorescent dye was applied to flowers of specific field margin plant species and NE sampled from within the bean crop and field margins using sweep-netting and pan-traps respectively. Captured insects were examined for the presence of the dye, indicative of a prior visit to the margin. Lady beetles and assassin bugs were most abundant in plots with B. pilosa margins; hoverflies with T. minuta and Parthenium hysterophorus margins; and lacewings with T. minuta and B. pilosa margins. Overall, NE benefitted from field margin plants, and those possessing extra floral nectaries had an added advantage. Field margin plants need careful selection to ensure benefits to different NE groups.

Floral nectar and pollen commonly contain diverse secondary metabolites. While these compounds are classically thought to play a role in plant defense, recent research indicates that they may also reduce disease in pollinators. Given that parasites have been implicated in ongoing bee declines, this discovery has spurred interest in the potential for 'medicinal' floral products to aid in pollinator conservation efforts. We review the evidence for antiparasitic effects of floral products on bee diseases, emphasizing the importance of investigating the mechanism underlying antiparasitic effects, including direct or host-mediated effects. We discuss the high specificity of antiparasitic effects of even very similar compounds, and highlight the need to consider how nonadditive effects of multiple compounds, and the post-ingestion transformation of metabolites, mediate the disease-reducing capacity of floral products. While the bulk of research on antiparasitic effects of floral products on bee parasites has been conducted in the lab, we review evidence for the impact of such effects in the field, and highlight areas for future research at the floral product-bee disease interface. Such research has great potential both to enhance our understanding of the role of parasites in shaping plant-bee interactions, and the role of plants in determining bee-parasite dynamics. This understanding may in turn reveal new avenues for pollinator conservation., Competing Interests: None., (© 2022 The Authors.)

Background: Hymenopteran parasitoids provide key natural pest regulation services and are reared commercially as biological control agents. Therefore, understanding parasitoid community composition in natural populations is important to enable better management for optimized natural pest regulation. We carried out a field study to understand the parasitoid community associated with Aphis fabae on East African smallholder farms. Either common bean (Phaseolus vulgaris) or lablab (Lablab purpureus) sentinel plants were infested with Aphis fabae and deployed in 96 fields across Kenya, Tanzania, and Malawi., Results: A total of 463 parasitoids emerged from sentinel plants of which 424 were identified by mitochondrial cytochrome oxidase I (COI) barcoding. Aphidius colemani was abundant in Kenya, Tanzania and Malawi, while Lysiphlebus testaceipes was only present in Malawi. The identity of Aphidius colemani specimens were confirmed by sequencing LWRh and 16S genes and was selected for further genetic and population analyses. A total of 12 Aphidius colemani haplotypes were identified. Of these, nine were from our East African specimens and three from the Barcode of Life Database (BOLD)., Conclusion: Aphidius colemani and Lysiphlebus testaceipes are potential targets for conservation biological control in tropical smallholder agro-ecosystems. We hypothesize that high genetic diversity in East African populations of Aphidius colemani suggests that this species originated in East Africa and has spread globally due to its use as a biological control agent. These East African populations could have potential for use as strains in commercial biological control or to improve existing Aphidius colemani strains by selective breeding., (© 2021 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.)

Caffeine is a widely occurring plant defense chemical 1 , 2 that occurs in the nectar of some plants, e.g., Coffea or Citrus spp., where it may influence pollinator behavior to enhance pollination. 3 , 4 Honey bees fed caffeine form longer lasting olfactory memory associations, 5 which could give plants with caffeinated nectar an adaptive advantage by inducing more visits to flowers. Caffeinated free-flying bees show enhanced learning performance 6 and are more likely to revisit a caffeinated target feeder or artificial flower, 7-9 although it is not clear whether improved memory of the target cues or the perception of caffeine as a reward is the cause. Here, we show that inexperienced bumble bees (Bombus terrestris) locate new food sources emitting a learned floral odor more consistently if they have been fed caffeine. In laboratory arena tests, we fed bees a caffeinated food alongside a floral odor blend (priming) and then used robotic experimental flowers 10 to disentangle the effects of caffeine improving memory for learned food-associated cues versus caffeine as a reward. Inexperienced bees primed with caffeine made more initial visits to target robotic flowers emitting the target odor compared to control bees or those primed with odor alone. Caffeine-primed bees tended to improve their floral handling time faster. Although the effects of caffeine were short lived, we show that food-locating behaviors in free-flying bumble bees can be enhanced by caffeine provided in the nest. Consequently, there is potential to redesign commercial colonies to enhance bees' forage focus or even bias bees to forage on a specific crop., Competing Interests: Declaration of interests F.L.W. is an employee of Biobest NV. F.L.W. was involved in the study design and interpretation, but the funding bodies themselves were not involved in the design, data collection, analysis, or decision to publish. At time of submission, the authors held no patents related to this work., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Background: Besides feeding on blood, females of the malaria vector Anopheles gambiae sensu lato readily feed on natural sources of plant sugars. The impact of toxic secondary phytochemicals contained in plant-derived sugars on mosquito physiology and the development of Plasmodium parasites remains elusive. The focus of this study was to explore the influence of the alkaloid ricinine, found in the nectar of the castor bean Ricinus communis, on the ability of mosquitoes to transmit Plasmodium falciparum., Methods: Females of Anopheles gambiae and its sibling species Anopheles coluzzii were exposed to ricinine through sugar feeding assays to assess the effect of this phytochemical on mosquito survival, level of P. falciparum infection and growth rate of the parasite., Results: Ricinine induced a significant reduction in the longevity of both Anopheles species. Ricinine caused acceleration in the parasite growth rate with an earlier invasion of the salivary glands in both species. At a concentration of 0.04 g l -1 in An. coluzzii, ricinine had no effect on mosquito infection, while 0.08 g l -1 ricinine-5% glucose solution induced a 14% increase in An. gambiae infection rate., Conclusions: Overall, our findings reveal that consumption of certain nectar phytochemicals can have unexpected and contrasting effects on key phenotypic traits that govern the intensity of malaria transmission. Further studies will be required before concluding on the putative role of ricinine as a novel control agent, including the development of ricinine-based toxic and transmission-blocking sugar baits. Testing other secondary phytochemicals in plant nectar will provide a broader understanding of the impact which plants can have on the transmission of vector-borne diseases., (© 2021. The Author(s).)

Common bean ( Phaseolus vulgaris ) production and storage are limited by numerous constraints. Insect pests are often the most destructive. However, resource-constrained smallholders in sub-Saharan Africa (SSA) often do little to manage pests. Where farmers do use a control strategy, it typically relies on chemical pesticides, which have adverse effects on the wildlife, crop pollinators, natural enemies, mammals, and the development of resistance by pests. Nature-based solutions -in particular, using biological control agents with sustainable approaches that include biopesticides, resistant varieties, and cultural tools-are alternatives to chemical control. However, significant barriers to their adoption in SSA include a lack of field data and knowledge on the natural enemies of pests, safety, efficacy, the spectrum of activities, the availability and costs of biopesticides, the lack of sources of resistance for different cultivars, and spatial and temporal inconsistencies for cultural methods. Here, we critically review the control options for bean pests, particularly the black bean aphid ( Aphis fabae ) and pod borers ( Maruca vitrata ). We identified natural pest regulation as the option with the greatest potential for this farming system. We recommend that farmers adapt to using biological control due to its compatibility with other sustainable approaches, such as cultural tools, resistant varieties, and biopesticides for effective management, especially in SSA.

Beneficial insect communities on farms are influenced by site- and landscape-level factors, with pollinator and natural enemy populations often associated with semi-natural habitat remnants. They provide ecosystem services essential for all agroecosystems. For smallholders, natural pest regulation may be the only affordable and available option to manage pests. We evaluated the beneficial insect community on smallholder bean farms (Phaseolus vulgaris L.) and its relationship with the plant communities in field margins, including margin trees that are not associated with forest fragments. Using traps, botanical surveys and transect walks, we analysed the relationship between the floral diversity/composition of naturally regenerating field margins, and the beneficial insect abundance/diversity on smallholder farms, and the relationship with crop yield. More flower visits by potential pollinators and increased natural enemy abundance measures in fields with higher plant, and particularly tree, species richness, and these fields also saw improved crop yields. Many of the flower visitors to beans and potential natural enemy guilds also made use of non-crop plants, including pesticidal and medicinal plant species. Selective encouragement of plants delivering multiple benefits to farms can contribute to an ecological intensification approach. However, caution must be employed, as many plants in these systems are introduced species., (© 2021. The Author(s).)

Emergent infectious diseases are one of the main drivers of species loss. Emergent infection with the microsporidian Nosema bombi has been implicated in the population and range declines of a suite of North American bumblebees, a group of important pollinators. Previous work has shown that phytochemicals found in pollen and nectar can negatively impact parasites in individuals, but how this relates to social epidemiology and by extension whether plants can be effectively used as pollinator disease management strategies remains unexplored. Here, we undertook a comprehensive screen of UK agri-environment scheme (AES) plants, a programme designed to benefit pollinators and wider biodiversity in agricultural settings, for phytochemicals in pollen and nectar using liquid chromatography and mass spectrometry. Caffeine, which occurs across a range of plant families, was identified in the nectar of sainfoin ( Onobrychis viciifolia ), a component of UK AES and a major global crop. We showed that caffeine significantly reduces N. bombi infection intensity, both prophylactically and therapeutically, in individual bumblebees ( Bombus terrestris ), and, for the first time, that such effects impact social epidemiology, with colonies reared from wild-caught queens having both lower prevalence and intensity of infection. Furthermore, infection prevalence was lower in foraging bumblebees from caffeine-treated colonies, suggesting a likely reduction in population-level transmission. Combined, these results show that N. bombi is less likely to be transmitted intracolonially when bumblebees consume naturally available caffeine, and that this may in turn reduce environmental prevalence. Consequently, our results demonstrate that floral phytochemicals at ecologically relevant concentrations can impact pollinator disease epidemiology and that planting strategies that increase floral abundance to support biodiversity could be co-opted as disease management tools.

Phytosterols are primary plant metabolites that have fundamental structural and regulatory functions. They are also essential nutrients for phytophagous insects, including pollinators, that cannot synthesize sterols. Despite the well-described composition and diversity in vegetative plant tissues, few studies have examined phytosterol diversity in pollen. We quantified 25 pollen phytosterols in 122 plant species (105 genera, 51 families) to determine their composition and diversity across plant taxa. We searched literature and databases for plant phylogeny, environmental conditions, and pollinator guilds of the species to examine the relationships with pollen sterols. 24-methylenecholesterol, sitosterol and isofucosterol were the most common and abundant pollen sterols. We found phylogenetic clustering of twelve individual sterols, total sterol content and sterol diversity, and of sterol groupings that reflect their underlying biosynthesis pathway (C-24 alkylation, ring B desaturation). Plants originating in tropical-like climates (higher mean annual temperature, lower temperature seasonality, higher precipitation in wettest quarter) were more likely to record higher pollen sterol content. However, pollen sterol composition and content showed no clear relationship with pollinator guilds. Our study is the first to show that pollen sterol diversity is phylogenetically clustered and that pollen sterol content may adapt to environmental conditions., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

Background: Many crops are dependent on pollination by insects. Habitat management in agricultural landscapes can support pollinator services and even augment crop production. Common bean ( Phaseolus vulgaris L.) is an important legume for the livelihoods of smallholder farmers in many low-income countries, particularly so in East Africa. While this crop is autogamous, it is frequently visited by pollinating insects that could improve yields. However, the value of pollination services to common beans (Kariasii) yield is not known., Methods: We carried out pollinator-exclusion experiments to determine the contribution of insect pollinators to bean yields. We also carried out a fluorescent-dye experiment to evaluate the role of field margins as refuge for flower-visitors., Results: Significantly higher yields, based on pods per plant and seeds per pod, were recorded from open -pollinated and hand -pollinated flowers compared to plants from which pollinators had been excluded indicating that flower visitors contribute significantly to bean yields. Similarly, open and hand -pollinated plants recorded the highest mean seed weight. Extrapolation of yield data to field scale indicated a potential increase per hectare from 681 kg in self -pollinated beans to 1,478 kg in open -pollinated beans indicating that flower visitors contributed significantly to crop yield of beans. Our marking study indicated that flower-visiting insects including bees, flies and lepidopterans moved from the field margin flowers into the bean crop. Overall, these results show that insect pollinators are important for optimising bean yields and an important food security consideration on smallholder farms. Field margin vegetation also provides habitat for flower-visiting insects that pollinate beans. Hence, non-crop habitats merit further research focusing on establishing which field margin species are most important and their capacity to support other ecosystem services such as natural pest regulation or even pests., Competing Interests: The authors declare there are no competing interests., (©2020 Elisante et al.)

Herbivory can induce chemical changes throughout plant tissues including flowers, which could affect pollinator-pathogen interactions. Pollen is highly defended compared to nectar, but no study has examined whether herbivory affects pollen chemistry. We assessed the effects of leaf herbivory on nectar and pollen alkaloids in Nicotiana tabacum, and how herbivory-induced changes in nectar and pollen affect pollinator-pathogen interactions. We damaged leaves of Nicotiana tabacum using the specialist herbivore Manduca sexta and compared nicotine and anabasine concentrations in nectar and pollen. We then pooled nectar and pollen by collection periods (within and after one month of flowering), fed them in separate experiments to bumble bees (Bombus impatiens) infected with the gut pathogen Crithidia bombi, and assessed infections after seven days. We did not detect alkaloids in nectar, and leaf damage did not alter the effect of nectar on Crithidia counts. In pollen, herbivory induced higher concentrations of anabasine but not nicotine, and alkaloid concentrations rose and then fell as a function of days since flowering. Bees fed pollen from damaged plants had Crithidia counts 15 times higher than bees fed pollen from undamaged plants, but only when pollen was collected after one month of flowering, indicating that both damage and time since flowering affected interaction outcomes. Within undamaged treatments, bees fed late-collected pollen had Crithidia counts 10 times lower than bees fed early-collected pollen, also indicating the importance of time since flowering. Our results emphasize the role of herbivores in shaping pollen chemistry, with consequences for interactions between pollinators and their pathogens.

Many pollinator species are declining due to a variety of interacting stressors including pathogens, sparking interest in understanding factors that could mitigate these outcomes. Diet can affect host-pathogen interactions by changing nutritional reserves or providing bioactive secondary chemicals. Recent work found that sunflower pollen (Helianthus annuus) dramatically reduced cell counts of the gut pathogen Crithidia bombi in bumble bee workers (Bombus impatiens), but the mechanism underlying this effect is unknown. Here we analyzed methanolic extracts of sunflower pollen by LC-MS and identified triscoumaroyl spermidines as the major secondary metabolite components, along with a flavonoid quercetin-3-O-hexoside and a quercetin-3-O-(6-O-malonyl)-hexoside. We then tested the effect of triscoumaroyl spermidine and rutin (as a proxy for quercetin glycosides) on Crithidia infection in B. impatiens, compared to buckwheat pollen (Fagopyrum esculentum) as a negative control and sunflower pollen as a positive control. In addition, we tested the effect of nine fatty acids from sunflower pollen individually and in combination using similar methods. Although sunflower pollen consistently reduced Crithidia relative to control pollen, none of the compounds we tested had significant effects. In addition, diet treatments did not affect mortality, or sucrose or pollen consumption. Thus, the mechanisms underlying the medicinal effect of sunflower are still unknown; future work could use bioactivity-guided fractionation to more efficiently target compounds of interest, and explore non-chemical mechanisms. Ultimately, identifying the mechanism underlying the effect of sunflower pollen on pathogens will open up new avenues for managing bee health.

Plants emit an extraordinary diversity of chemicals that provide information about their identity and mediate their interactions with insects. However, most studies of this have focused on a few model species in controlled environments, limiting our capacity to understand plant-insect chemical communication in ecological communities. Here, by integrating information theory with ecological and evolutionary theories, we show that a stable information structure of plant volatile organic compounds (VOCs) can emerge from a conflicting information process between plants and herbivores. We corroborate this information "arms race" theory with field data recording plant-VOC associations and plant-herbivore interactions in a tropical dry forest. We reveal that plant VOC redundancy and herbivore specialization can be explained by a conflicting information transfer. Information-based communication approaches can increase our understanding of species interactions across trophic levels., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Besides providing food and shelter to natural enemies of crop pests, plants used in conservation biological control interventions potentially provide additional ecosystem services including providing botanical insecticides. Here we concurrently tested the strength of these two services from six non-crop plants in managing cabbage pests in Ghana over three successive field seasons. Crop margin plantings of Ageratum conyzoides, Tridax procumbens, Crotalaria juncea, Cymbopogon citratus, Lantana camara and Talinum triangulare were compared with a bare earth control in a three-way split plot design such that the crop in each plot was sprayed with either a 10% (w/v) aqueous extract from the border plant species, a negative control (water) and a positive control (emamectin benzoate 'Attack' insecticide). Pests were significantly less numerous in all unsprayed treatments with non-crop plant margins and in corresponding sprayed treatments (with botanical or synthetic insecticide positive control) while treatments with bare earth margin or sprayed with water (negative controls) had the highest pest densities. Numbers of predators were significantly depressed by synthetic insecticide but higher in other treatments whether unsprayed or sprayed with botanical insecticide. We conclude that some plant species have utility in both conservation biological control and as source of botanical insecticides that are relatively benign to natural enemies. In this crop system, however, the additional cost associated with using botanical insecticides was not justified by greater levels of pest suppression than achieved from border plants alone.

Immune systems provide a key defence against diseases. However, they are not a panacea and so both vertebrates and invertebrates co-opt naturally occurring bioactive compounds to treat themselves against parasites and pathogens. In vertebrates, this co-option is complex, with pharmacodynamics leading to differential effects of treatment at different life stages, which may reflect age-linked differences in the immune system. However, our understanding of pharmacodynamics in invertebrates is almost non-existent. Critically, this knowledge may elucidate broad parallels across animals in regard to the requirement for the co-option of bioactive compounds to ameliorate disease. Here, we used biochanin A, an isoflavone found in the pollen of red clover ( Trifolium pratense ), to therapeutically treat Nosema bombi (Microsporidia) infection in bumblebee ( Bombus terrestris ) larvae and adults, and thus examine age-linked pharmacodynamics in an invertebrate. Therapeutic treatment of larvae with biochanin A did not reduce the infection intensity of N. bombi in adults. In contrast, therapeutic treatment of adults did reduce the infection intensity of N. bombi This transition in parasite resistance to bioactive compounds mirrors the age-linked pharmacodynamics of vertebrates. Understanding how different life-history stages respond to therapeutic compounds will provide novel insights into the evolution of foraging and self-medication behaviour in natural systems more broadly., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Fall Armyworm (FAW) ( Spodoptera frugiperda ) is a polyphagous and highly destructive pest of many crops. It was recently introduced into Africa and now represents a serious threat to food security, particularly because of yield losses in maize, which is the staple food for the majority of small-scale farmers in Africa. The pest has also led to increased production costs, and threatens trade because of quarantines imposed on produce from the affected countries. There is limited specific knowledge on its management among smallholders since it is such a new pest in Africa. Some synthetic insecticides have been shown to be effective in controlling FAW, but in addition to the economic, health and environmental challenges of pesticide use insecticide resistance is highly prevalent owing to years of FAW management in the Americas. Therefore, there is a need for the development and use of alternatives for the management of FAW. These include plant-derived pesticides. Here we review the efficacy and potential of 69 plant species, which have been evaluated against FAW, and identify opportunities for use among small-scale maize farmers with a focus on how pesticidal plants might be adopted in Africa for management of FAW. The biological activities were diverse and included insecticidal, insectistatic (causing increased larval duration), larvicidal, reduced growth and acute toxicity (resulting in adverse effects within a short time after exposure). While most of these studies have been conducted on American plant taxa many South American plants are now cosmopolitan weeds so these studies are relevant to the African context.