Multiple stressors: using the honeybee model BEEHAVE to explore how spatial and temporal forage stress affects colony resilience

The causes underlying the increased mortality of honeybee colonies (Apis mellifera) observed over the past decade remain unclear. Since so far the evidence for monocausal explanations is equivocal, involvement of multiple stressors is generally assumed. We here focus on various aspects of forage availability, which have received less attention than other stressors because it is virtually impossible to explore them empirically. We applied the colony model BEEHAVE, which links within-hive dynamics and foraging, to stylized landscape settings to explore how foraging distance, forage supply, and “forage gaps”, i.e. periods in which honeybees cannot find any nectar and pollen, affect colony resilience and the mechanisms behind. We found that colony extinction was mainly driven by foraging distance, but the timing of forage gaps had strongest effects on time to extinction. Sensitivity to forage gaps of 15 days was highest in June or July even if otherwise forage availability was sufficient to survive. Forage availability affected colonies via cascading effects on queen's egg-laying rate, reduction of new-emerging brood stages developing into adult workers, pollen debt, lack of workforce for nursing, and reduced foraging activity. Forage gaps in July led to reduction in egg-laying and increased mortality of brood stages at a time when the queen's seasonal egg-laying rate is at its maximum, leading to colony failure over time. Our results demonstrate that badly timed forage gaps interacting with poor overall forage supply reduce honeybee colony resilience. Existing regulation mechanisms which in principle enable colonies to cope with varying forage supply in a given landscape and year, such as a reduction in egg-laying, have only a certain capacity. Our results are hypothetical, as they are obtained from simplified landscape settings, but they are consistent with existing empirical knowledge. They offer ample opportunities for testing the predicted effects of forage stress in controlled experiments

REVIEW: Towards a systems approach for understanding honeybee decline: a stocktaking and synthesis of existing models

Published© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.Summary 1. The health of managed and wild honeybee colonies appears to have declined substantially in Europe and the United States over the last decade. Sustainability of honeybee colonies is important not only for honey production, but also for pollination of crops and wild plants alongside other insect pollinators. A combination of causal factors, including parasites, pathogens, land use changes and pesticide usage, are cited as responsible for the increased colony mortality. 2. However, despite detailed knowledge of the behaviour of honeybees and their colonies, there are no suitable tools to explore the resilience mechanisms of this complex system under stress. Empirically testing all combinations of stressors in a systematic fashion is not feasible. We therefore suggest a cross-level systems approach, based on mechanistic modelling, to investigate the impacts of (and interactions between) colony and land management. 3. We review existing honeybee models that are relevant to examining the effects of different stressors on colony growth and survival. Most of these models describe honeybee colony dynamics, foraging behaviour or honeybee – varroa mite – virus interactions. 4. We found that many, but not all, processes within honeybee colonies, epidemiology and foraging are well understood and described in the models, but there is no model that couples in-hive dynamics and pathology with foraging dynamics in realistic landscapes. 5. Synthesis and applications. We describe how a new integrated model could be built to simulate multifactorial impacts on the honeybee colony system, using building blocks from the reviewed models. The development of such a tool would not only highlight empirical research priorities but also provide an important forecasting tool for policy makers and beekeepers, and we list examples of relevant applications to bee disease and landscape management decisions.Biotechnology and Biological Sciences Research Council (BBSRC

Searching for nests of the invasive Asian hornet (Vespa velutina) using radio-telemetry

This is the final version of the article. Available from the publisher via the DOI in this recordAsian hornets (Vespa velutina) are voracious predators of bees, and are the latest emerging threat to managed and wild pollinator populations in Europe. To prevent establishment or reduce the rate of spread of V. velutina, early detection and destruction of nests is considered the only option. Detection is difficult as their nests are well hidden and flying hornets are difficult to follow over long distances. We address this challenge by tracking individual V. velutina workers flying back to their nests using radio telemetry for the first time, finding five previously undiscovered nests, up to 1.33 km from hornet release points. Hornets can fly with 0.28 g tags if the tag:hornet ratio is less than 0.8. This method offers a step-change in options to tackle the spread of this invader, providing an efficient means of finding V. velutina nests in complex environments to manage this emerging threat to pollinators.We thank Olivier Bonnard for discussions, sourcing materials, and advice on locating foraging V. velutina workers at INRA Bordeaux-Aquitaine. We also thank members of Jersey Beekeeping Association for their assistance in catching V. velutina workers in Jersey. The work was funded by a Defra research project grant (PH0532), with additional support by the States of Jersey Department of Environment and generous philanthropic donations by the South West Beekeeping Associations’ Forum (SWBKF), Somerset Beekeeping Association, Dorset Beekeeping Association, Cornwall Beekeeping Association, West Cornwall Beekeeping Association, Devon Beekeeping Association, Bournemouth & South Dorset Beekeeping Association, and B.J. Sherriff. We are grateful to INRA Bordeaux-Aquitaine, States of Jersey Department of Environment, and Durrell Wildlife Park for their welcome and permission to use facilities at their institutions

Predicting honeybee colony failure: using the BEEHAVE model to simulate colony responses to pesticides

PublishedJournal ArticleResearch Support, Non-U.S. Gov'tTo simulate effects of pesticides on different honeybee (Apis mellifera L.) life stages, we used the BEEHAVE model to explore how increased mortalities of larvae, in-hive workers, and foragers, as well as reduced egg-laying rate, could impact colony dynamics over multiple years. Stresses were applied for 30 days, both as multiples of the modeled control mortality and as set percentage daily mortalities to assess the sensitivity of the modeled colony both to small fluctuations in mortality and periods of low to very high daily mortality. These stresses simulate stylized exposure of the different life stages to nectar and pollen contaminated with pesticide for 30 days. Increasing adult bee mortality had a much greater impact on colony survival than mortality of bee larvae or reduction in egg laying rate. Importantly, the seasonal timing of the imposed mortality affected the magnitude of the impact at colony level. In line with the LD50, we propose a new index of "lethal imposed stress": the LIS50 which indicates the level of stress on individuals that results in 50% colony mortality. This (or any LISx) is a comparative index for exploring the effects of different stressors at colony level in model simulations. While colony failure is not an acceptable protection goal, this index could be used to inform the setting of future regulatory protection goals.J.R. was funded to do this work on an Industrial CASE PhD studentship funded by the Biology and Biotechnology Sciences Research Council of the UK (BBSRC), and Syngenta. J.O., M.B., and P.K. were supported on BBSRC project BB/K014463/

A review of the success of the UK strategy to tackle the invasive insect Vespa velutina nigrithorax, the “Asian hornet”

Interreg Atlantic Area project: Atlantic Positive – Conservation of Atlantic Pollination services and control of the invasive species Vespa velutina. Work-package 8; Action No. 1: Strategy to prevent the spread of V. velutina to the UK. To prevent Vespa velutina from spreading across the British mainland, avoiding economic loss and harm to biodiversity, this report describes action plans and specific measures for the British mainland and the maritime space between UK and mainland Europe.The UK’s strategy to prevent the establishment of the Asian hornet, Vespa velutina nigrithorax, and/or manage its presence and impacts is embedded within the UK’s generic strategy for invasive non-native species, but specifically informed by the non-native species risk assessment for Vespa velutina and the pest-specific contingency plan for the Asian hornet. This report introduces relevant UK policies and legislation for invasive non-native species, before summarising the UK’s assessment of risk of entry, establishment and impact, and its rapid response plans addressing the Asian hornet’s periodic incursions in the UK. The report describes the effectiveness of those plans to manage and control the spread of the Asian hornet on the British mainland over the last seven years; and provides a reflection on the next steps required to ensure establishment is delayed as long as possible, while preparing for a time when the species may become established in the UK. The details may provide useful pointers for other European countries in which the Asian hornet is not yet established, but also incorporates suggestions for the UK based on actions from regions where the hornet has already become established.European Regional Development Fund (ERDF) Interreg Atlantic Area ProgrammeBiotechnology and Biological Sciences Research Council (BBSRC

BEEHAVE: A systems model of honeybee colony dynamics and foraging to explore multifactorial causes of colony failure

Journal Article© 2014 The Authors. Journal of Applied Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly citedSummary: A notable increase in failure of managed European honeybee Apis mellifera L. colonies has been reported in various regions in recent years. Although the underlying causes remain unclear, it is likely that a combination of stressors act together, particularly varroa mites and other pathogens, forage availability and potentially pesticides. It is experimentally challenging to address causality at the colony scale when multiple factors interact. In silico experiments offer a fast and cost-effective way to begin to address these challenges and inform experiments. However, none of the published bee models combine colony dynamics with foraging patterns and varroa dynamics. We have developed a honeybee model, BEEHAVE, which integrates colony dynamics, population dynamics of the varroa mite, epidemiology of varroa-transmitted viruses and allows foragers in an agent-based foraging model to collect food from a representation of a spatially explicit landscape. We describe the model, which is freely available online (www.beehave-model.net). Extensive sensitivity analyses and tests illustrate the model's robustness and realism. Simulation experiments with various combinations of stressors demonstrate, in simplified landscape settings, the model's potential: predicting colony dynamics and potential losses with and without varroa mites under different foraging conditions and under pesticide application. We also show how mitigation measures can be tested. Synthesis and applications. BEEHAVE offers a valuable tool for researchers to design and focus field experiments, for regulators to explore the relative importance of stressors to devise management and policy advice and for beekeepers to understand and predict varroa dynamics and effects of management interventions. We expect that scientists and stakeholders will find a variety of applications for BEEHAVE, stimulating further model development and the possible inclusion of other stressors of potential importance to honeybee colony dynamics. © 2014 The Authors. Journal of Applied Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.Biotechnology and Biological Sciences Research Council (BBSRC

On the structure of buoyant fires with varying levels of fuel-turbulence

This paper employs a novel burner to study the effects of fuel-generated turbulence on the spatial and temporal structure of buoyant turbulent diffusion flames which are representative of large fires. Fuel-turbulence levels are increased using a perforated plate that issues high-velocity jets, enabling shearing of the fuel stream. The perforated plate may be recessed to control the turbulence level at the jet exit plane. It is shown that the exit plane axial velocity fluctuations can be increased from 0.135 m/s to 1.813 m/s. Varying the levels of fuel-turbulence in the burner allows for the control of key processes defining buoyant fires such as the spatial and temporal flame structure and flame instability modes. These processes are characterised by high-speed simultaneous imaging of planar laser-induced fluorescence of the OH radical (OH-PLIF) and Mie scattering from soot particles. Increasing the fuel-turbulence level deforms the flame, which promotes non-radial lateral entrainment into the flame sheet. This results in a sharp increase in the tilting of the near-field flame sheet along the vertical flame axis. Strong angular entrainment forces are shown to overcome the diffusive and thermal expansive forces at the flame neck, which leads to a strained asymmetric sinuous flame pinch-off instability, followed by separation of the flame base. Sinuous pinch-off instabilities occur at a greater frequency than the symmetric varicose pinch-off instabilities observed for flames with low fuel-turbulence. The asymmetric stretching of the flame neck inhibits the formation of the classical puffing instability formed with an axisymmetric plume that defines classically buoyant flames. Probability density functions calculated for the flame front curvature and flame surface area are shown to monotonically broaden in the near-field region of the flame due to lateral entrainment effects. The transition to buoyancy-driven turbulence also shifts to an increasingly more upstream location. This burner, with its well-defined boundary conditions and novel data, forms a platform for advancing capabilities to model complex fire phenomena including turbulence-buoyancy interactions

Statin Use in Relation to Intraocular Pressure, Glaucoma, and Ocular Coherence Tomography Parameters in the UK Biobank

PURPOSE. The purpose of this study was to evaluate the relationship between statin use and glaucoma-related traits. METHODS. In a cross-sectional study, we included 118,153 UK Biobank participants with data on statin use and corneal-compensated IOP. In addition, we included 192,283 participants (8982 cases) with data on glaucoma status. After excluding participants with neurodegenerative diseases, 41,638 participants with macular retinal nerve fiber layer thickness (mRNFL) and 41,547 participants with macular ganglion cell inner plexiform layer thickness (mGCIPL) were available for analysis. We examined associations of statin use with IOP, mRNFL, mGCIPL, and glaucoma status utilizing multivariable-adjusted regression models. We assessed whether a glaucoma polygenic risk score (PRS) modified associations. We performed Mendelian randomization (MR) experiments to investigate associations with various glaucoma-related outcomes. RESULTS. Statin users had higher unadjusted mean IOP ± SD than nonusers, but in a multivariable-adjusted model, IOP did not differ by statin use (difference = 0.05 mm Hg, 95% confidence interval [CI] = −0.02 to 0.13, P = 0.17). Similarly, statin use was not associated with prevalent glaucoma (odds ratio [OR] = 1.05, 95% CI = 0.98 to 1.13). Statin use was weakly associated with thinner mRNFL (difference = −0.15 microns, 95% CI = −0.28 to −0.01, P = 0.03) but not with mGCIPL thickness (difference = −0.12 microns, 95% CI = −0.29 to 0.05, P = 0.17). No association was modified by the glaucoma PRS (Pinteraction ≥ 0.16). MR experiments showed no evidence for a causal association between the cholesterol-altering effect of statins and several glaucoma traits (inverse weighted variance P ≥ 0.14). CONCLUSIONS. We found no evidence of a protective association between statin use and glaucoma or related traits after adjusting for key confounders

Matrix permeability of reservoir rocks, Ngatamariki geothermal field, Taupo Volcanic Zone, New Zealand

The Taupo Volcanic Zone (TVZ) hosts 23 geothermal fields, seven of which are currently utilised for power generation. Ngatamariki geothermal field (NGF) is one of the latest geothermal power generation developments in New Zealand (commissioned in 2013), located approximately 15 km north of Taupo. Samples of reservoir rocks were taken from the Tahorakuri Formation and Ngatamariki Intrusive Complex, from five wells at the NGF at depths ranging from 1354 to 3284 m. The samples were categorised according to whether their microstructure was pore or microfracture dominated. Image analysis of thin sections impregnated with an epoxy fluorescent dye was used to characterise and quantify the porosity structures and their physical properties were measured in the laboratory. Our results show that the physical properties of the samples correspond to the relative dominance of microfractures compared to pores. Microfracture-dominated samples have low connected porosity and permeability, and the permeability decreases sharply in response to increasing confining pressure. The pore-dominated samples have high connected porosity and permeability, and lower permeability decrease in response to increasing confining pressure. Samples with both microfractures and pores have a wide range of porosity and relatively high permeability that is moderately sensitive to confining pressure. A general trend of decreasing connected porosity and permeability associated with increasing dry bulk density and sonic velocity occurs with depth; however, variations in these parameters are more closely related to changes in lithology and processes such as dissolution and secondary veining and re-crystallisation. This study provides the first broad matrix permeability characterisation of rocks from depth at Ngatamariki, providing inputs for modelling of the geothermal system. We conclude that the complex response of permeability to confining pressure is in part due to the intricate dissolution, veining, and recrystallization textures of many of these rocks that lead to a wide variety of pore shapes and sizes. While the laboratory results are relevant only to similar rocks in the Taupo Volcanic Zone, the relationships they highlight are applicable to other geothermal fields, as well as rock mechanic applications to, for example, aspects of volcanology, landslide stabilisation, mining, and tunnelling at depth