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Optimal nutrition is crucial for honey bee colony growth and robust immune systems. Honey bee nutrition is complex and depends on the floral composition of the landscape. Foraging behavior of honey bees depends on both colony environment and external environment. There are significant gaps in knowledge regarding honey bee nutrition, and hence no optimal diet is available for honey bees, as there is for other livestock. In this review, we discuss (1) foraging behavior of honey bees, (2) nutritional needs, (3) nutritional supplements used by beekeepers, (4) probiotics, and (5) supplemental forage and efforts integrating floral diversity into cropping systems.

Although poor nutrition is cited as one of the crucial factors in global pollinator decline, the requirements and role of several important nutrients (especially micronutrients) in honey bees are not well understood. Micronutrients, viz. phytosterols, play a physiologically vital role in insects as precursors of important molting hormones and building blocks of cellular membranes. There is a gap in comprehensive understanding of the impacts of dietary sterols on honey bee physiology. In the present study, we investigated the role of 24-methylenecholesterol—a key phytosterol—in honey bee nutritional physiology. Artificial diets with varying concentrations of 24-methylenecholesterol (0%, 0.1%. 0.25%, 0.5%, 0.75%, and 1% dry diet weight) were formulated and fed to honey bees in a laboratory cage experiment. Survival, diet consumption, head protein content, and abdominal lipid contents were significantly higher in dietary sterol-supplemented bees. Our findings provide additional insights regarding the role of this important sterol in honey bee nutritional physiology. The insights gleaned from this study could also advance the understanding of sterol metabolism and regulation in other bee species that are dependent on pollen for sterols, and assist in formulation of a more complete artificial diet for honey bees (Apis mellifera Linnaeus, 1758) (Hymenoptera: Apidae).

Honey bee larvae are dependent on the social structure of colony for their provisioning and survival. With thousands of larvae being managed collectively by groups of foragers (collecting food resources) and nurse bees (processing food and provisioning larvae), coordination of colony efforts in rearing brood depends on multiple dynamic cues of larval presence and needs. Much of these cues appear to be chemical, with larvae producing multiple pheromones, major being brood ester pheromone (BEP; nonvolatile blend of fatty acid esters) that elicits both short-term releaser effects and long-term primer effects. While BEP can affect colony food collection and processing with the signaling of larval presence, it is unclear if BEP signals individual larval needs. To understand this aspect, in a series of experiments we manipulated larval feeding environment by depriving larvae from adult bee contact for 4-h period and examined (1) nurse bee interactions with contact-deprived and nondeprived larvae and larval extracts; (2) forager bee responses to contact-deprived and nondeprived larval extracts. We also characterized BEP of contact-deprived and nondeprived larvae. We found that nurse honey bees tend to aggregate more over contact-deprived larvae when compared with nondeprived larvae, but that these effects were not found in response to whole hexane extracts. Our analytical results suggest that BEP components changed in both quantity and quality over short period of contact deprivation. These changes affected foraging behavior, but did not appear to directly affect nursing behavior, suggesting that different chemical cues are involved in regulating nursing effort to individual larvae.

Researchers often collect and analyze corbicular pollen from honey bees to identify the plant sources on which they forage for pollen or to estimate pesticide exposure of bees via pollen. Described herein is an effective pollen-trapping method for collecting corbicular pollen from honey bees returning to their hives. This collection method results in large quantities of corbicular pollen that can be used for research purposes. Honey bees collect pollen from many plant species, but typically visit one species during each collection trip. Therefore, each corbicular pollen pellet predominantly represents one plant species, and each pollen pellet can be described by color. This allows the sorting of samples of corbicular pollen by color to segregate plant sources. Researchers can further classify corbicular pollen by analyzing the morphology of acetolyzed pollen grains for taxonomic identification. These methods are commonly used in studies related to pollinators such as pollination efficiency, pollinator foraging dynamics, diet quality, and diversity. Detailed methodologies are presented for collecting corbicular pollen using pollen traps, sorting pollen by color, and acetolyzing pollen grains. Also presented are results pertaining to the frequency of pellet colors and taxa of corbicular pollen collected from honey bees in five different cropping systems.

Simple Summary Phytosterols are important micronutrients that are essential for production of insect molting hormones and cellular membrane integrity. Past research has shown that the key phytosterol that honey bees need is 24-methylenecholesterol. This phytosterol improves honey bee longevity and sustains brood production. Hence, it is important to understand how 24-methylenecholesterol can shape honey bee physiology by altering protein profiles of vital honey bee tissues. Nurse bees secrete glandular secretions (brood food) using hypopharyngeal and mandibular glands in their head regions. Further, it has been shown that this sterol is selectively accumulated in nurse bee heads. Thus, it is imperative to examine the protein profiles of nurse bee heads, in response to dietary 24-methylenecholesterol manipulation. In this study, groups of newly emerged nurse bees were fed with varying concentrations of dietary 24-methylenecholesterol, while the control groups received no sterol. We found that dietary sterol manipulation altered the protein profiles in nurse bee heads, with important nutritional marker proteins being upregulated in high dietary sterol groups. The important proteins identified in this study may serve as vital markers of nutritional stress related to sterols in honey bees, paving the way for future research on bee nutrition. Abstract Phytosterols are important micronutrients that are precursors of important molting hormones and help maintain cellular membrane integrity in insects including bees. Previous research has shown that 24-methylenecholesterol is a key phytosterol that enhances honey bee longevity and improves nurse bee physiology. Nurse bees have the ability to selectively transfer this sterol to developing larvae through brood food. This study examines the physiological impacts of 24-methylenecholesterol on nurse bees, by analyzing the protein profiles of nurse bee heads upon dietary sterol manipulation. Dietary experimental groups consisting of newly emerged honey bees were provided with varying concentrations of 24-methylenecholesterol for three weeks. At the end of the study, honey bees were collected and proteomic analysis was performed on honey bee heads. A total of 1715 proteins were identified across experimental groups. The mean relative abundances of nutritional marker proteins (viz. major royal jelly proteins 1, 4, 5, 7) were higher in experimental groups supplemented with higher dietary sterol concentrations, when compared with the control dietary group. The mean relative abundances of important enzymatic proteins (aminopeptidase and calcium-transporting ATPase) were higher in control groups, whereas mean relative abundances of oxysterol-binding protein and fatty acid-binding protein were higher in higher dietary sterol groups.

Global decline in insect pollinators, especially bees, have resulted in extensive research into understanding the various causative factors and formulating mitigative strategies. For commercial beekeepers in the United States, overwintering honey bee colony losses are significant, requiring tactics to overwinter bees in conditions designed to minimize such losses. This is especially important as overwintered honey bees are responsible for colony expansion each spring, and overwintered bees must survive in sufficient numbers to nurse the spring brood and forage until the new ‘replacement’ workers become fully functional. In this study, we examined the physiology of overwintered (diutinus) bees following various overwintering storage conditions. Important physiological markers, i.e., head proteins and abdominal lipid contents were higher in honey bees that overwintered in controlled indoor storage facilities, compared with bees held outdoors through the winter months. Our findings provide new insights into the physiology of honey bees overwintered in indoor and outdoor environments and have implications for improved beekeeping management.

Honey bees provision glandular secretions in the form of royal jelly as larval nourishment to developing queens. Exposure to chemicals and nutritional conditions can influence queen development and thus impact colony fitness. Previous research reports that royal jelly remains pesticide-free during colony-level exposure and that chemical residues are buffered by the nurse bees. However, the impacts of pesticides can also manifest in quality and quantity of royal jelly produced by nurse bees. Here, we tested how colony exposure to a multi-pesticide pollen treatment influences the amount of royal jelly provisioned per queen and the additional impacts on royal jelly nutritional quality. We observed differences in the metabolome, proteome, and phytosterol compositions of royal jelly synthesized by nurse bees from multi-pesticide exposed colonies, including significant reductions of key nutrients such as 24-methylenecholesterol, major royal jelly proteins, and 10-hydroxy-2-decenoic acid. Additionally, quantity of royal jelly provisioned per queen was lower in colonies exposed to pesticides, but this effect was colony-dependent. Pesticide treatment had a greater impact on royal jelly nutritional composition than the weight of royal jelly provisioned per queen cell. These novel findings highlight the indirect effects of pesticide exposure on queen developmental nutrition and allude to social consequences of nurse bee glandular degeneration.

Poor nutrition is an important factor in global bee population declines. A significant gap in knowledge persists regarding the role of various nutrients (especially micronutrients) in honey bees. Sterols are essential micronutrients in insect diets and play a physiologically vital role as precursors of important molting hormones and building blocks of cellular membranes. Sterol requirements and metabolism in honey bees are poorly understood. Among all pollen sterols, 24-methylenecholesterol is considered the key phytosterol required by honey bees. Nurse bees assimilate this sterol from dietary sources and store it in their tissues as endogenous sterol, to be transferred to the growing larvae through brood food. This study examined the duration of replacement of such endogenous sterols in honey bees. The dietary 13C-labeled isotopomer of 24-methylenecholesterol added to artificial bee diet showed differential, progressive in vivo assimilation across various honey bee tissues. Significantly higher survival, diet consumption, head protein content and abdominal lipid content were observed in the dietary sterol-supplemented group than in the control group. These findings provide novel insights into phytosterol utilization and temporal pattern of endogenous 24-methylenecholesterol replacement in honey bees.

The decline in populations of insect pollinators is a global concern. While multiple factors are implicated, there is uncertainty surrounding the contribution of certain groups of pesticides to losses in wild and managed bees. Nanotechnology-based pesticides (NBPs) are formulations based on multiple particle sizes and types. By packaging active ingredients in engineered particles, NBPs offer many benefits and novel functions, but may also exhibit different properties in the environment when compared with older pesticide formulations. These new properties raise questions about the environmental disposition and fate of NBPs and their exposure to pollinators. Pollinators such as honey bees have evolved structural adaptations to collect pollen, but also inadvertently gather other types of environmental particles which may accumulate in hive materials. Knowledge of the interaction between pollinators, NBPs, and other types of particles is needed to better understand their exposure to pesticides, and essential for characterizing risk from diverse environmental contaminants. The present review discusses the properties, benefits and types of nanotechnology-based pesticides, the propensity of bees to collect such particles and potential impacts on bee pollinators.

Background Significant annual honey bee colony losses have been reported in the USA and across the world over the past years. Malnutrition is one among several causative factors for such declines. Optimal nutrition serves as the first line of defense against multiple stressors such as parasites/pathogens and pesticides. Given the importance of nutrition, it is imperative to understand bee nutrition holistically, identifying dietary sources that may fulfill bee nutritional needs. Pollen is the primary source of protein for bees and is critical for brood rearing and colony growth. Currently, there is significant gap in knowledge regarding the chemical and nutritional composition of pollen. Methods Targeted sterol analysis and untargeted metabolomics were conducted on five commercially available crop pollens, three bee-collected crop pollens, three vegetable oils (often added to artificial protein supplements by beekeepers), and one commonly used artificial protein supplement. Results This study reports key phytosterols and metabolites present across a spectrum of bee diets, including some of the major bee-pollinated crop pollens in the western United States. Significant differences were observed in sterol concentrations among the dietary sources tested. Among all quantified sterols, the highest concentrations were observed for 24-methylenecholesterol and further, pollen samples exhibited the highest 24-methylenecholesterol among all diet sources that were tested. Also, 236 metabolites were identified across all dietary sources examined. Conclusion Information gleaned from this study is crucial in understanding the nutritional landscape available to all bee pollinators and may further assist in future efforts to develop comprehensive database of nutrients and metabolites present in all bee diets.

Pesticide stress is one of the important factors for global bee declines. Apart from physiological and developmental anomalies, pesticides also impose cognitive damages on bees. The present study investigates the visual acuity of wild populations of honey bees, in an agricultural intensification landscape, and corroborates the findings with controlled laboratory experiments. Even though overall morphometric examinations revealed no significant differences between the populations, correct color choices by bees in pesticide exposed populations were significantly reduced. The study reports, for the first time, the significant reduction in ommatidia facet diameter in these populations, as viewed under scanning electron microscope, along with the molecular underpinnings to these findings. Western blot studies revealed a significant reduction in expression of two visual proteins – blue-sensitive opsin and rhodopsin – in the pesticide exposed populations in both field and laboratory conditions. The novel findings from this study form the basis for further investigations into the effects of field realistic doses of multiple pesticide exposures on wild populations of honey bees.

Global western honey bee, Apis mellifera (L.) (Hymenoptera: Apidae), colony declines pose a significant threat to food production worldwide. Poor nutrition resulting from habitat loss, extensive monocultures, and agricultural intensification is among the several suggested drivers for colony declines. Pollen is the primary source of protein for honey bees; therefore, both pollen abundance and diversity are critical for colony growth and survival. Many cropping systems that employ honey bee colonies for pollination may lack sufficient pollen diversity and abundance to provide optimal bee nutrition. In this observational study, we documented the diversity and relative abundance of pollen collected by honey bees in five major pollinator-dependent crops in the western United States. We sampled pollen from pollen traps installed on honey bee colonies in the following cropping systems—almond, cherry, highbush blueberry, hybrid carrot, and meadowfoam. The pollen diversity was estimated by documenting the number of different pollen pellet colors and plant taxa found in each pollen sample. The lowest pollen diversity was found in almond crop. Relatively higher quantities of pollen collection were collected in almond, cherry, and meadowfoam cropping systems. The information gleaned from this study regarding pollen diversity and abundance may help growers, land managers, and beekeepers improve pollen forage available to bees in these cropping systems.

Pesticide exposures can have detrimental impacts on bee pollinators, ranging from immediate mortality to sub-lethal impacts. Flupyradifurone is the active ingredient in Sivanto™ and sulfoxaflor is the active ingredient in Transform®. They are both relatively new insecticides developed with an intent to reduce negative effects on bees, when applied to bee-attractive crops. With the growing concern regarding pollinator health and pollinator declines, it is important to have a better understanding of any potential negative impacts, especially sub-lethal, of these pesticides on bees. This study reports novel findings regarding physiological stress experienced by bees exposed to field application rates of these two insecticides via a Potter Tower sprayer. Two contact exposure experiments were conducted-a shorter 6-hour study and a longer 10-day study. Honey bee mortality, sugar syrup and water consumption, and physiological responses (oxidative stress and apoptotic protein assays) were assessed in bees exposed to Sivanto™ and Transform®, and compared to bees in control group. For the longer, 10-day contact exposure experiment, only the Sivanto™ group was compared to the control group, as high mortality recorded in the sulfoxaflor treatment group during the shorter contact exposure experiment, made the latter group unfeasible to test in the longer 10-days experiment. In both the studies, sugar syrup and water consumptions were significantly different between treatment groups and controls. The highest mortality was observed in Transform® exposed bees, followed by the Sivanto™ exposed bees. Estimates of reactive oxygen/nitrogen species indicated significantly elevated oxidative stress in both pesticide treatment groups, when compared to controls. Caspase-3 protein assays, an indicator of onset of apoptosis, was also significantly higher in the pesticide treatment groups. These differences were largely driven by post exposure duration, indicating sub-lethal impacts. Further, our findings also emphasize the need to revisit contact exposure impacts of Sivanto™, given the sub-lethal impacts and mortality observed in our long-term (10-day) contact exposure experiment.

In honey bees and many other social insects, production of queens is a vital task, as colony fitness is dependent on queens. The factors considered by honey bee workers in selecting larvae to rear new queens during emergency queen rearing are poorly understood. Identifying these parameters is critical, both in an evolutionary and apicultural context. As female caste development in honey bees is dependent on larval diet (i.e. nutrition), we hypothesized that larval nutritional state is meticulously assessed and used by workers in selection of larvae for queen rearing. To test this hypothesis, we conducted a series of experiments manipulating the nutritional status of one day old larvae by depriving them of brood food for a four-hour period, and then allowing workers to choose larvae for rearing queens from nutritionally deprived and non-deprived larvae. We simultaneously investigated the role of genetic relatedness in selection of larvae for queen rearing. In all the experiments, significantly greater numbers of non-deprived larvae than deprived larvae were selected for queen rearing irrespective of genetic relatedness. Our results demonstrate that honey bees perceive the nutritional state of larvae and use that information when selecting larvae for rearing queens in the natural emergency queen replacement process.

Pesticides have been reported to be one of the major drivers in the global pollinator losses. The large-scale decline in honey bees, an important pollinator group, has resulted in comprehensive studies on honey bee colonies. Lack of information on native wild pollinators has paved the way for this study, which highlights the underlying evolutionary changes occurring in the wild honey bee populations exposed to pesticides along an agricultural intensification landscape. The study reports an increased genetic diversity in native Apis cerana Fabricius (Hymenoptera: Apidae) populations continually exposed to pesticide stress. An increased heterozygosity, evidenced by a higher electrophoretic banding pattern, was observed in the pesticide-exposed populations for two isozymes involved with xenobiotic metabolism—esterase and glucose-6-phosphate dehydrogenase. Differential banding patterns also revealed a higher percentage of polymorphic loci, number of polymorphic bands, Nei’s genetic distance, etc. observed in these populations in the Randomly Amplified Polymorphic DNA–Polymerase Chain Reaction (RAPD–PCR) experiments using three random decamer primers. Higher heterozygosity, being indicative of a more resistant population, implies population survival within the threshold pesticide stress. This study reports such changes for the first time in native wild Indian honey bee populations exposed to pesticides and has far-reaching implications on the population adaptability under pesticide stress.

Little information is available regarding the adverse effects of pesticides on natural honey bee populations. This study highlights the detrimental effects of pesticides on honey bee olfaction through behavioural studies, scanning electron microscopic imaging of antennal sensillae and confocal microscopic studies of honey bee brains for calcium ions on Apis cerana, a native Indian honey bee species. There was a significant decrease in proboscis extension response and biologically active free calcium ions and adverse changes in antennal sensillae in pesticide exposed field honey bee populations compared to morphometrically similar honey bees sampled from low/no pesticide sites. Controlled laboratory experiments corroborated these findings. This study reports for the first time the changes in antennal sensillae, expression of Calpain 1(an important calcium binding protein) and resting state free calcium in brains of honey bees exposed to pesticide stress.

International audience; AbstractPesticides have been cited as one of the major drivers of pollinator loss. However, little is known about pesticide impacts on natural populations of native honey bee species. This study looked into the effect of pesticides with respect to oxidative stress in the laboratory and in field populations of two native Indian honey bee species (Apis dorsata and A. cerana) by examining a combination of biomarkers, e.g., superoxide dismutase, catalase and xanthine oxidase. A significant upregulation of all three biomarkers was observed in both treated individuals in laboratory experiments and field populations sampled from a pesticide use gradient. This study reports, for the first time, an increase in expression of xanthine oxidase in an invertebrate system (honey bees) exposed to pesticides.