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Eusocial insects live in teeming societies with thousands of their kin. In this crowded environment, workers combat disease by removing or burying their dead or diseased nestmates. For honey bees, we found that hygienic brood-removal behavior is triggered by two odorants – β-ocimene and oleic acid – which are released from brood upon freeze-killing. β-ocimene is a co-opted pheromone that normally signals larval food-begging, whereas oleic acid is a conserved necromone across arthropod taxa. Interestingly, the odorant blend can induce hygienic behavior more consistently than either odorant alone. We suggest that the volatile β-ocimene flags hygienic workers’ attention, while oleic acid is the death cue, triggering removal. Bees with high hygienicity detect and remove brood with these odorants faster than bees with low hygienicity, and both molecules are strong ligands for hygienic behavior-associated odorant binding proteins (OBP16 and OBP18). Odorants that induce low levels of hygienic behavior, however, are weak ligands for these OBPs. We are therefore beginning to paint a picture of the molecular mechanism behind this complex behavior, using odorants associated with freeze-killed brood as a model.

The honey bee is responsible for pollination of a large proportion of crop plants, but the health of honey bee populations has been challenged by the parasitic mite Varroa destructor . Mite infestation is the main cause of colony losses during the winter months, which causes significant economic challenges in apiculture. Treatments have been developed to control the spread of varroa. However, many of these treatments are no longer effective due to acaricide resistance. In a search of varroa-active compounds, we tested the effect of dialkoxybenzenes on the mite. A structure–activity relationship revealed that 1-allyloxy-4-propoxybenzene is most active of a series of dialkoxybenzenes tested. We found that three compounds (1-allyloxy-4-propoxybenzene, 1,4-diallyloxybenzene and 1,4-dipropoxybenzene) cause paralysis and death of adult varroa mites, whereas the previously discovered compound, 1,3-diethoxybenzene, which alters host choice of adult mites in certain conditions, did not cause paralysis. Since paralysis can be caused by inhibition of acetylcholinesterase (AChE), a ubiquitous enzyme in the nervous system of animals, we tested dialkoxybenzenes on human, honey bee and varroa AChE. These tests revealed that 1-allyloxy-4-propoxybenzene had no effects on AChE, which leads us to conclude that 1-allyloxy-4-propoxybenzene does not exert its paralytic effect on mites through AChE. In addition to paralysis, the most active compounds affected the ability of the mites to find and remain at the abdomen of host bees provided during assays. A test of 1-allyloxy-4-propoxybenzene in the field, during the autumn of 2019 in two locations, showed that this compound has promise in the treatment of varroa infestations.

Honey bee viruses are serious pathogens that can cause poor colony health and productivity. We analyzed a multi-year longitudinal dataset of abundances of nine honey bee viruses (deformed wing virus A, deformed wing virus B, black queen cell virus, sacbrood virus, Lake Sinai virus, Kashmir bee virus, acute bee paralysis virus, chronic bee paralysis virus, and Israeli acute paralysis virus) in colonies located across Canada to describe broad trends in virus intensity and occurrence among regions and years. We also tested climatic variables (temperature, wind speed, and precipitation) as predictors in an effort to understand possible drivers underlying seasonal patterns in viral prevalence. Temperature was a significant positive predictor of the total number of viruses per sample, which was highest in British Columbia (mean = 5.0). Lake Sinai virus (LSV) was the most prevalent overall (at 89%) and had the highest infection intensity, at an average of 3.9 × 10 8 copies per bee. Acute bee paralysis virus was the least prevalent virus (at 4.7%) and had the lowest infection intensity (1.9 × 10 5 copies per bee). Surprisingly, including Varroa abundance as a covariate did not significantly improve model fit for any virus. All viruses, except Kashmir bee virus, varied by region, and one or more climatic variables were significant predictors for six of the nine viruses. Although climatic effects were often inconsistent among individual viruses, we show that climatic variables can be better predictors of virus intensity and occurrence than Varroa mite abundance, at least when infestation rates are low.

We present a novel way to select for highly polygenic traits. For millennia, humans have used observable phenotypes to selectively breed stronger or more productive livestock and crops. Selection on genotype, using single-nucleotide polymorphisms (SNPs) and genome profiling, is also now applied broadly in livestock breeding programs; however, selection on protein/peptide or mRNA expression markers has not yet been proven useful. Here we demonstrate the utility of protein markers to select for disease-resistant hygienic behavior in the European honey bee ( Apis mellifera L.). Robust, mechanistically-linked protein expression markers, by integrating cis- and trans- effects from many genomic loci, may overcome limitations of genomic markers to allow for selection. After three generations of selection, the resulting marker-selected stock outperformed an unselected benchmark stock in terms of hygienic behavior, and had improved survival when challenged with a bacterial disease or a parasitic mite, similar to bees selected using a phenotype–based assessment for this trait. This is the first demonstration of the efficacy of protein markers for industrial selective breeding in any agricultural species, plant or animal.

The honey bee queen produces pheromones that function in both releaser and primer roles such as attracting a retinue of workers around her, attracting drones on mating flights, preventing workers from reproducing at the individual (worker egg-laying) and colony (swarming) level, and regulating several other aspects of colony functioning. The queen mandibular pheromone (QMP), consisting of five synergistic components, is the only pheromone chemically identified in the honey bee (Apis mellifera L.) queen, but this pheromone does not fully duplicate the pheromonal activity of a full queen extract. To identify the remaining unknown compounds for retinue attraction, honey bee colonies were selectively bred to have low response to synthetic QMP and high response to a queen extract in a laboratory retinue bioassay. Workers from these colonies were then used in the bioassay to guide the isolation and identification of the remaining active components. Four new compounds were identified from several glandular sources that account for the majority of the difference in retinue attraction between synthetic QMP and queen extract: methyl (Z)-octadec-9-enoate (methyl oleate), (E)-3-(4-hydroxy-3-methoxyphenyl)-prop-2-en-1-ol (coniferyl alcohol), hexadecan-1-ol, and (Z9,Z12,Z15)-octadeca-9,12,15-trienoic acid (linolenic acid). These compounds were inactive alone or in combination, and they only elicited attraction in the presence of QMP. There was still unidentified activity remaining in the queen extract. The queen therefore produces a synergistic, multiglandular pheromone blend of at least nine compounds for retinue attraction, the most complex pheromone blend known for inducing a single behavior in any organism.

We examined the effect of larval and adult nutrition on worker honey bee (Apis mellifera L.) ovary development. Workers were fed high or low-pollen diets as larvae, and high or low-protein diets as adults. Workers fed low-protein diets at both life stages had the lowest levels of ovary development, followed by those fed high-protein diets as larvae and low- quality diets as adults, and then those fed diets poor in protein as larvae but high as adults. Workers fed high-protein diets at both life stages had the highest levels of ovary development. The increases in ovary development due to improved dietary protein in the larval and adult life stages were additive. Adult diet also had an effect on body mass. The results demonstrate that both carry-over of larval reserves and nutrients acquired in the adult life stage are important to ovary development in worker honey bees. Carry-over from larval development, however, appears to be less important to adult fecundity than is adult nutrition. Seasonal trends in worker ovary development and mass were examined throughout the brood rearing season. Worker ovary development was lowest in spring, highest in mid-summer, and intermediate in fall.

We assessed bee diversity and abundance in urban areas of Vancouver, British Columbia, Canada, to determine how urban environments can support bees. Habitats examined were community and botanical gardens, urban wild areas, Naturescape flower beds and backyards, and traditional flower beds and backyards. A total of 56 bee species (Hymenoptera), including species of the genera Andrena Fabr. (Andrenidae), Bombus Latr. (Apidae), Osmia Panzer and Megachile Latr. (Megachilidae), and Halictus Latr. and Dialictus Pauly (Halictidae), were collected. Abundance exhibited strong seasonal variation. Wild bees were most abundant during late spring, whereas honey bees peaked at the end of the summer. The most abundant species seen was the managed honey bee Apis mellifera L. (Apidae), followed by wild Bombus flavifrons Cresson. Community and botanical gardens, and plants such as cotoneaster (Cotoneaster Medik. sp.) and blackberry (Rubus discolor Weihe & Nees) (Rosaceae), centaurea (Centaurea L. sp.; Asteraceae), buttercup (Ranunculus L. sp.; Ranunculaceae), and foxglove (Digitalis L. sp.; Scrophulariaceae), had the highest abundance of bees, while bee populations in wild areas were the most diverse. Weeds such as dandelions (Taraxacum officinale G.H. Weber ex Wiggers; Asteraceae) dominated these wild areas and had one of the highest diversities of bee visitors. Traditional flower beds with tulips (Tulipa L. sp.; Liliaceae) and petunias (Petunia Juss. sp.; Solanaceae) had relatively poor diversity and abundance of bees throughout the year. Our study suggests that urban areas have the potential to be important pollinator reservoirs, especially if both bloom and habitat heterogeneity are maintained and enhanced through sustainable urban planning. Nous avons estimé la diversité et l'abondance d'abeilles dans différents biotopes urbains de la ville de Vancouver, Colombie-Britannique, Canada, afin de déterminer comment les environnements urbains peuvent supporter les abeilles. Les habitats examinés incluent les jardins potagers et botaniques, les environs urbains sauvages, les parterres et les arrière-cours dits Naturescape, et les parterres et les arrière-cours traditionnels. Nous avons ressemblé un total de 56 espèces d'abeilles (Hymenoptera) dont Andrena Fabr. (Andrenidae), Bombus Latr. (Apidae), Osmia Panzer et Megachile Latr. (Megachilidae), Halictus Latr. et Dialictus Pauly (Halictidae). L'abondance a présenté une forte variation saisonnière. Les abeilles féeriques étaient considérablement plus abondantes pendant la dernière moitié du printemps, tandis que les abeilles d'élevage étaient plus abondantes pendant la dernière moitié de l'été. L'espèce la plus abondante était l'abeille d'élevage, Apis mellifera L. (Apidae), suivie de l'espèce féerique Bombus flavifrons Cresson. Les jardins potagers et botaniques des plantes telles que Cotoneaster Medik sp. et Rubus discolor Wehie & Ness (Rosaceae), Centaurea L. sp. (Asteraceae), Ranunculus L. sp. (Ranunculaceae), et Digitalis L. sp. (Scrophulariaceae) ont présenté la plus élevée abondance d'abeilles alors que les environs urbains sauvages ont exhibé les plus diverses populations d'abeilles. « Mauvaise herbes », comme les Taraxacum officinale G.H. Weber ex Wiggers (Asteraceae), dominaient ces habitats et présentaient la diversité d'abeilles visiteuses las plus élevée. Les parterres traditionnels, avec fleures telles que les Tulipa L. sp. (Liliaceae) et les Petunia Juss. sp. (Solanaceae), présentaient une diversité d'abeilles relativement pauvre tout au cour de l'an. Notre étude suggère que les zones urbaines ont du potentiel comme important réservoir de pollinisation, surtout quand et l'hétérogénéité d'habitat et hétérogénéité de fleures sont maintenues et accrues à travers le développement urbain durable.

Abstract British Columbia beekeepers, like many beekeepers around the world, are currently facing declines in honey bee health and high overwinter colony losses. To better understand the economics and the cycle of yearly colony loss and replacement of this critical agricultural industry, we collected and analyzed survey data on beekeeping costs and returns. Forty British Columbia beekeepers provided details about revenue sources, variable costs, capital costs, and investments. Ten surveyed beekeepers managed between 1 and 9 colonies, 10 managed between 10 and 39 colonies, 9 managed between 40 and 100 colonies, 5 managed between 101 and 299 colonies, 3 managed between 300 and 699 colonies, and 3 managed 700 colonies or more. The data was used to calculate beekeeping profit and to parameterize a model that explores the economic impact of colony loss rates and replacement strategies. Survey results show that when the data is aggregated, revenues exceed costs for beekeeping operations in British Columbia with a per colony profit of $56.92 or $0.87 per pound of honey produced. Surveyed operations with fewer than 100 colonies have negative profits, while operations with 100–299 colonies have positive profits. Surveyed operations in the Cariboo, North Coast, and Okanagan regions have the highest profits while surveyed operations in the Peace region have the lowest profits. Profit modeling shows that replacing losses with packages generates lower profit than replacing losses with split colonies. Our modeling shows that operations that diversify their revenue to include bee sales and commercial pollination accrue higher profits and can withstand higher winter loss rates.

We studied the abundance, diversity, and dispersion patterns of managed and wild bee (Hymenoptera: Apoidea) populations in commercial highbush blueberry and cranberry (Ericaceae: Vaccinium corymbosum L., Vaccinium macrocarpon Ait.) fields in the Fraser Valley of British Columbia, and assessed their potential as pollinators of these crops by determining which groups of bees had the greatest impact on percent yield and mass of berries. Bumble bees were evenly distributed within both crops. Other wild bee species were well distributed in blueberry fields but generally remained at edges of cranberry fields. Percent berry yield was not related to bee abundance for any group of bees, nor was species diversity correlated with berry mass. Blueberry mass and cranberry mass were related to abundance of bumble bees but not to that of honey bees or other wild bees. Bumble bees are recommended as potential alternative pollinators of these crops. Nous avons étudié l’abondance, la diversité et la distribution d’abeilles mellifères et sauvages (Hymenoptera: Apoidea) dans des champs de bleuets (« highbush ») et de canneberges (Ericaceae: Vaccinium corymbosum L., Vaccinium macrocarpon Ait.) au sein de la vallée Fraser en Colombie Britannique, et nous avons évalué leur potentiel en tant que pollinisateurs en déterminant quels groupes d’abeilles avaient le plus d’impact sur les récoltes et la masse des baies. Les bourdons se sont bien répartis dans les deux types de culture. Les autres abeilles sauvages se sont aussi bien réparties dans les champs de bleuets mais sont restées le plus souvent en périphérie des champs de canneberges. Le pourcentage des récoltes de baies n’était pas relié à l’abondance d’aucun groupe d’abeille. Aucune corrélation n’a été trouvé entre la diversité des abeilles et la masse des baies. Nous avons parcontre trouvé que la masse des bleuets et des canneberges était relié à l’abondance des bourdons mais non à celle des abeilles mellifères ou autres abeilles sauvages. Les bourdons sont donc recommandés comme pollinisateurs potentiels et alternatifs pour ces deux types de cultures.

Highbush blueberry pollination depends on managed honey bees (Apis mellifera) L. for adequate fruit sets; however, beekeepers have raised concerns about the poor health of colonies after pollinating this crop. Postulated causes include agrochemical exposure, nutritional deficits, and interactions with parasites and pathogens, particularly Melisococcus plutonius [(ex. White) Bailey and Collins, Lactobacillales: Enterococcaceae], the causal agent of European foulbrood disease, but other pathogens could be involved. To broadly investigate common honey bee pathogens in relation to blueberry pollination, we sampled adult honey bees from colonies at time points corresponding to before (t1), during (t2), at the end (t3), and after (t4) highbush blueberry pollination in British Columbia, Canada, across 2 years (2020 and 2021). Nine viruses, as well as M. plutonius, Vairimorpha ceranae, and V. apis [Tokarev et al., Microsporidia: Nosematidae; formerly Nosema ceranae (Fries et al.) and N. apis (Zander)], were detected by PCR and compared among colonies located near and far from blueberry fields. We found a significant interactive effect of time and blueberry proximity on the multivariate pathogen community, mainly due to differences at t4 (corresponding to ∼6 wk after the beginning of the pollination period). Post hoc comparisons of pathogens in near and far groups at t4 showed that detections of sacbrood virus (SBV), which was significantly higher in the near group, not M. plutonius, was the primary driver. Further research is needed to determine if the association of SBV with highbush blueberry pollination is contributing to the health decline that beekeepers observe after pollinating this crop.