Honey bee‐collected pollen in agro‐ecosystems reveals diet diversity,diet quality,and pesticide exposure

European honey bees Apis mellifera are important commercial pollinators that have suffered greater than normal overwintering losses since 2007 in North America and Europe. Contributing factors likely include a combination of parasites, pesticides, and poor nutrition. We examined diet diversity, diet nutritional quality, and pesticides in honey bee‐collected pollen from commercial colonies in the Canadian Maritime Provinces in spring and summer 2011. We sampled pollen collected by honey bees at colonies in four site types: apple orchards, blueberry fields, cranberry bogs, and fallow fields. Proportion of honey bee‐collected pollen from crop versus noncrop flowers was high in apple, very low in blueberry, and low in cranberry sites. Pollen nutritional value tended to be relatively good from apple and cranberry sites and poor from blueberry and fallow sites. Floral surveys ranked, from highest to lowest in diversity, fallow, cranberry, apple, and blueberry sites. Pesticide diversity in honey bee‐collected pollen was high from apple and blueberry sites and low from cranberry and fallow sites. Four different neonicotinoid pesticides were detected, but neither these nor any other pesticides were at or above LD₅₀ levels. Pollen hazard quotients were highest in apple and blueberry sites and lowest in fallow sites. Pollen hazard quotients were also negatively correlated with the number of flower taxa detected in surveys. Results reveal differences among site types in diet diversity, diet quality, and pesticide exposure that are informative for improving honey bee and land agro‐ecosystem management.     

Differential sensitivity of honey bees and bumble bees to a dietary insecticide (imidacloprid)

Currently, there is concern about declining bee populations and the sustainability of pollination services. One potential threat to bees is the unintended impact of systemic insecticides, which are ingested by bees in the nectar and pollen from flowers of treated crops. To establish whether imidacloprid, a systemic neonicotinoid and insect neurotoxin, harms individual bees when ingested at environmentally realistic levels, we exposed adult worker bumble bees, Bombus terrestris L. (Hymenoptera: Apidae), and honey bees, Apis mellifera L. (Hymenoptera: Apidae), to dietary imidacloprid in feeder syrup at dosages between 0.08 and 125 μg l^?1. Honey bees showed no response to dietary imidacloprid on any variable that we measured (feeding, locomotion and longevity). In contrast, bumble bees progressively developed over time a dose-dependent reduction in feeding rate with declines of 10–30% in the environmentally relevant range of up to 10 μg l^?1, but neither their locomotory activity nor longevity varied with diet. To explain their differential sensitivity, we speculate that honey bees are better pre-adapted than bumble bees to feed on nectars containing synthetic alkaloids, such as imidacloprid, by virtue of their ancestral adaptation to tropical nectars in which natural alkaloids are prevalent. We emphasise that our study does not suggest that honey bee colonies are invulnerable to dietary imidacloprid under field conditions, but our findings do raise new concern about the impact of agricultural neonicotinoids on wild bumble bee populations.     

Comparative study on the dynamics and performances of Apis mellifera jemenitica and imported hybrid honeybee colonies in southwestern Saudi Arabia

The aims of this study were to assess the seasonal population dynamics and evaluate the performance of Apis mellifera jemenitica (local bee) and introduced hybrid honeybee colonies in the lowlands and highlands of southwestern Saudi Arabia. Data regarding the performance and population dynamics parameters such as brood and adult bee population, amounts of stored pollen and nectar were gathered from the two races (25 colonies of each) for one year (April 2013 through March 2014), and statistically tested. The results indicated that at low lands, local bee colonies maintained relatively high brood and adult bee populations (P < 0.05) than introduced honeybee colonies and produced more (P < 0.05) honey. The local bee colonies were able to hoard three times more (P < 0.05) pollen and built more (P < 0.05) queen cells than introduced bees in both the low and highland areas. The annual survival rate of local bee colonies was almost double (P < 0.05) than that of introduced honeybee colonies. Moreover, local bees had greater (P < 0.05) adult bee and brood populations than imported, throughout the year. The relatively good performance of local colonies could be due to their long year’s adaptation to cope with resource scarcity and unpredictable environmental conditions of the regions. The possible reasons for the dwindling of the imported hybrid colonies could be due to continuing to exhibit adaptive characteristics of their original that might not fit well with the new environment.     

Changes in nutritional composition from bee pollen to pollen patty used in bumblebee rearing

Pollen patties are generally supplied to augment the nourishment of honey bee or bumblebee colonies during late winter or early spring. In the present study, we examined nutrient content of bee collected pollen (BP) and pollen patties (PT) prepared from those bee collected pollens in order to understand the chemical compositional change of pollen to pollen patty and figure out the possible benefits for bee’s health. Protein and fat contents of pollen patties were found lower than that reported for the respective bee pollens. Amino and fatty acids followed the same trend for the simple logical reason. In contrast, carbohydrate contents of pollen patties were found much higher than the bee pollen. The addition of sugar solution in the process of pollen patty preparation could presumably explain the change. Another possible determinant factor of bee’s preference of feed protein to fat ratio was found the highest for PT 2 (5.5) followed by PT 1 and BP 2 (4.1), BP Ref. (2.9) and the least value obtained for BP 1 (2.2). Additionally, we discuss the possible health benefits if the bee collected pollen is used as a human food supplement.     

Effects of a neonicotinoid pesticide on thermoregulation of African honey bees (Apis mellifera scutellata)

Thiamethoxam is a widely used neonicotinoid pesticide that, as agonist of the nicotinic acetylcholine receptors, has been shown to elicit a variety of sublethal effects in honey bees. However, information concerning neonicotinoid effects on honey bee thermoregulation is lacking. Thermoregulation is an essential ability for the honey bee that guarantees the success of foraging and many in-hive tasks, especially brood rearing. We tested the effects of acute exposure to thiamethoxam (0.2, 1, 2 ng/bee) on the thorax temperatures of foragers exposed to low (22 °C) and high (33 °C) temperature environments. Thiamethoxam significantly altered honey bee thorax temperature at all doses tested; the effects elicited varied depending on the environmental temperature and pesticide dose to which individuals were exposed. When bees were exposed to the high temperature environment, the high dose of thiamethoxam increased their thorax temperature 1–2 h after exposure. When bees were exposed to the low temperature, the higher doses of the neonicotinoid reduced bee thorax temperatures 60–90 min after treatment. In both experiments, the neonicotinoid decreased the temperature of bees the day following the exposure. After a cold shock (5 min at 4 °C), the two higher doses elicited a decrease of the thorax temperature, while the lower dose caused an increase, compared to the control. These alterations in thermoregulation caused by thiamethoxam may affect bee foraging activity and a variety of in-hive tasks, likely leading to negative consequences at the colony level. Our results shed light on sublethal effect of pesticides which our bees have to deal with.     

The neonicotinoids thiacloprid,imidacloprid, and clothianidin affect the immunocompetence of honey bees (Apis mellifera L.)

A strong immune defense is vital for honey bee health and colony survival. This defense can be weakened by environmental factors that may render honey bees more vulnerable to parasites and pathogens. Honey bees are frequently exposed to neonicotinoid pesticides, which are being discussed as one of the stress factors that may lead to colony failure. We investigated the sublethal effects of the neonicotinoids thiacloprid, imidacloprid, and clothianidin on individual immunity, by studying three major aspects of immunocompetence in worker bees: total hemocyte number, encapsulation response, and antimicrobial activity of the hemolymph. In laboratory experiments, we found a strong impact of all three neonicotinoids. Thiacloprid (24 h oral exposure, 200 μg/l or 2000 μg/l) and imidacloprid (1 μg/l or 10 μg/l) reduced hemocyte density, encapsulation response, and antimicrobial activity even at field realistic concentrations. Clothianidin had an effect on these immune parameters only at higher than field realistic concentrations (50–200 μg/l). These results suggest that neonicotinoids affect the individual immunocompetence of honey bees, possibly leading to an impaired disease resistance capacity.     

Effects of pollen dilution on infection of Nosema ceranae in honey bees

Multiple stressors are currently threatening honey bee health, including pests and pathogens. Among honey bee pathogens, Nosema ceranae is a microsporidian found parasitizing the western honey bee (Apis mellifera) relatively recently. Honey bee colonies are fed pollen or protein substitute during pollen dearth to boost colony growth and immunity against pests and pathogens. Here we hypothesize that N. ceranae intensity and prevalence will be low in bees receiving high pollen diets, and that honey bees on high pollen diets will have higher survival and/or increased longevity. To test this hypothesis we examined the effects of different quantities of pollen on (a) the intensity and prevalence of N. ceranae and (b) longevity and nutritional physiology of bees inoculated with N. ceranae. Significantly higher spore intensities were observed in treatments that received higher pollen quantities (1:0 and 1:1 pollen:cellulose) when compared to treatments that received relatively lower pollen quantities. There were no significant differences in N. ceranae prevalence among different pollen diet treatments. Interestingly, the bees in higher pollen quantity treatments also had significantly higher survival despite higher intensities of N. ceranae. Significantly higher hypopharyngeal gland protein was observed in the control (no Nosema infection, and receiving a diet of 1:0 pollen:cellulose), followed by 1:0 pollen:cellulose treatment that was inoculated with N. ceranae. Here we demonstrate that diet with higher pollen quantity increases N. ceranae intensity, but also enhances the survival or longevity of honey bees. The information from this study could potentially help beekeepers formulate appropriate protein feeding regimens for their colonies to mitigate N. ceranae problems.     

Fatty acid and nutritive quality of chia (Salvia hispanica L.) seeds and plant during growth

The fatty acid (FA) profile, chemical composition, gross energy and organic matter digestibility of chia (Salvia hispanica L.) have been determined in the seed and in the plant collected at five progressive morphological stages from early vegetative to budding stage. The FA analyses disclosed quantitative differences between the plant stages that were characterised by a high percentage of polyunsaturated fatty acids (PUFA), which made up from 752 to 623 g/kg of the total FA of the plant during the growth cycle. The α-linolenic acid (ALA, C_18:3n–3) decreased from 649 g/kg, at the early vegetative stage, to 499 g/kg of the total FA, at the budding stage, while all the other FAs increased with increasing growth stage. The chia seed FAs were also highly unsaturated, with their main components being ALA (641 g/kg of the total FA) and linoleic acid (LA, C_18:2n–6; 188 g/kg of the total FA).The evolution of the quality of chia is closely related to the ageing of the plant. The chia plant provides a forage with a good nutritive value when harvested at a stage before the shooting period. After this, the nutritional quality of the plant considerably decreases with an increase in the fibrous fractions and a dramatical decrease of the crude protein content.     

Sperm viability and gene expression in honey bee queens (Apis mellifera) following exposure to the neonicotinoid insecticide imidacloprid and the organophosphate acaricide coumaphos

Honey bee population declines are of global concern. Numerous factors appear to cause these declines including parasites, pathogens, malnutrition and pesticides. Residues of the organophosphate acaricide coumaphos and the neonicotinoid insecticide imidacloprid, widely used to combat Varroa mites and for crop protection in agriculture, respectively, have been detected in wax, pollen and comb samples. Here, we assess the effects of these compounds at different doses on the viability of sperm stored in the honey bee queens’ spermatheca. Our results demonstrate that sub-lethal doses of imidacloprid (0.02 ppm) decreased sperm viability by 50%, 7 days after treatment. Sperm viability was a downward trend (about 33%) in queens treated with high doses of coumaphos (100 ppm), but there was not significant difference. The expression of genes that are involved in development, immune responses and detoxification in honey bee queens and workers exposed to chemicals was measured by qPCR analysis. The data showed that expression levels of specific genes were triggered 1 day after treatment. The expression levels of P450 subfamily genes, CYP306A1, CYP4G11 and CYP6AS14 were decreased in honey bee queens treated with low doses of coumaphos (5 ppm) and imidacloprid (0.02 ppm). Moreover, these two compounds suppressed the expression of genes related to antioxidation, immunity and development in queens at day 1. Up-regulation of antioxidants by these compounds in worker bees was observed at day 1. Coumaphos also caused a repression of CYP306A1 and CYP4G11 in workers. Antioxidants appear to prevent chemical damage to honey bees. We also found that DWV replication increased in workers treated with imidacloprid. This research clearly demonstrates that chemical exposure can affect sperm viability in queen honey bees.     

Crop Pollination Exposes Honey Bees to Pesticides Which Alters Their Susceptibility to the Gut Pathogen Nosema ceranae

Recent declines in honey bee populations and increasing demand for insect-pollinated crops raise concerns about pollinator shortages. Pesticide exposure and pathogens may interact to have strong negative effects on managed honey bee colonies. Such findings are of great concern given the large numbers and high levels of pesticides found in honey bee colonies. Thus it is crucial to determine how field-relevant combinations and loads of pesticides affect bee health. We collected pollen from bee hives in seven major crops to determine 1) what types of pesticides bees are exposed to when rented for pollination of various crops and 2) how field-relevant pesticide blends affect bees’ susceptibility to the gut parasite Nosema ceranae. Our samples represent pollen collected by foragers for use by the colony, and do not necessarily indicate foragers’ roles as pollinators. In blueberry, cranberry, cucumber, pumpkin and watermelon bees collected pollen almost exclusively from weeds and wildflowers during our sampling. Thus more attention must be paid to how honey bees are exposed to pesticides outside of the field in which they are placed. We detected 35 different pesticides in the sampled pollen, and found high fungicide loads. The insecticides esfenvalerate and phosmet were at a concentration higher than their median lethal dose in at least one pollen sample. While fungicides are typically seen as fairly safe for honey bees, we found an increased probability of Nosema infection in bees that consumed pollen with a higher fungicide load. Our results highlight a need for research on sub-lethal effects of fungicides and other chemicals that bees placed in an agricultural setting are exposed to.     

Nutritional effects of supplementary diets on brood development,biological activities and honey production of Apis mellifera L

The present research work was conducted to assess the impact of nutrient-enriched diet on the physiological activities and subsequently honey yield. Eighteen colonies of Apis mellifera L. were selected from Dera Ismail Khan region, KPK, Pakistan, during the winter and summer seasons, 2019–2020. Five pollen supplement diets were prepared and provided to screen out the palatable one to be fed as pollen alternative nutrition to bee bread. Results of diet consumption regarding mean data for consumption rate displayed that soybean flour enriched artificial diet was maximally consumed (74.34 g) by honey bees per week. Minimum consumption was observed for grinded groundnut enriched diet (64.62 g) which was relatively lesser than the other tested artificial diets. Results of area of worker brood disclosed that soybean flour fortified diet (1489.27 cm²/colony) statistically noteworthy than the other artificial diets whereas control (463.51 cm²/colony) was least effective. Highest bee strength (10.00 bee frames/colony) was noted in the bee colonies fed with soybean flour fortified diet, date paste (8.0 bee frames/colony) was the next effective one, among the tested pollen replacement diets whereas relatively least (5 bee frames/colony) was noticed in case of grinded groundnut. Highest body weight (12.41 g) of neonate bees was noted in case of soybean enriched diet while lowermost (5.31 g) was noted in control bees. Results of wax cell built up and foraging efficiency were also superior in artificial diets than respective control bees. Hence, artificial diets especially soybean-enriched pollen alternative diet can boost up the physiology of honey bee leading to increased honey yield and profit.     

Floral biology of the saguaro (Cereus giganteus)

A saguaro cactus (Cereus giganteus) produces an average of 295 flowers per season, each of which produces 286 mg fresh weight of pollen and 543 mg of nectar containing 24% sugar. At 7600 pollen grains/mg pollen, the yearly output per saguaro plant is 6.4×10⁸ grains. Based on the measured saguaro density of 6.56 plants/ha, 553 g/ha of pollen is produced yearly. The enormous variation among individual plants in terms of flower numbers and floral bloom patterns is documented.Honey bees (Apis mellifera L.), the main collectors of saguaro pollen, collect an average of 12.2 mg pollen per foraging trip and can thus harvest 23.5 pollen loads from one flower. An average honey bee colony collects 290 g of saguaro pollen over the season, which is 24.4% of their total intake. Individual colonies exhibit wide variation in pollen collecting activities with some closely tracking the pollen resource and others almost totally ignoring it. The average for seven colonies indicates that even though variation is great the overall trend is toward closely tracking and exploiting the saguaro pollen resource. Based on the pollen productivity of saguaro and a hypothetical 90% pollen harvesting efficiency of bees, the pollen harvest potential of the saguaro environment is 1.72 colony equivalents of pollen/ha and 0.5/ha for saguaro alone. This is the first quantitative reporting of the total pollen productivity and pollen resource utilization for any plant and an opportunistic pollinator.     

Effects of brood pheromone (SuperBoost) on consumption of protein supplement and growth of honey bee (Hymenoptera: Apidae) colonies during fall in a northern temperate climate

Honey bee, Apis mellifera L. (Hymenoptera: Apidae), nutrition is vital for colony growth and maintenance of a robust immune system. Brood rearing in honey bee colonies is highly dependent on protein availability. Beekeepers in general provide protein supplement to colonies during periods of pollen dearth. Honey bee brood pheromone is a blend of methyl and ethyl fatty acid esters extractable from cuticle of honey bee larvae that communicates the presence of larvae in a colony. Honey bee brood pheromone has been shown to increase protein supplement consumption and growth of honey bee colonies in a subtropical winter climate. Here, we tested the hypothesis that synthetic brood pheromone (SuperBoost) has the potential to increase protein supplement consumption during fall in a temperate climate and thus increase colony growth. The experiments were conducted in two locations in Oregon during September and October 2009. In both the experiments, colonies receiving brood pheromone treatment consumed significantly higher protein supplement and had greater brood area and adult bees than controls. Results from this study suggest that synthetic brood pheromone may be used to stimulate honey bee colony growth by stimulating protein supplement consumption during fall in a northern temperate climate, when majority of the beekeepers feed protein supplement to their colonies.     

Behavioral responses of the small hive beetle,Aethina tumida,to odors of three meliponine bee species and honey bees,Apis mellifera scutellata

Recent studies have shown that honey bees, bumble bees, and some meliponine bee species of the genera Trigona, Meliponula, and Dactylurina are hosts of the small hive beetle (SHB) Aethina tumidaMurray (Coleoptera: Nitidulidae), a pest of honey bee colonies in various regions of the world. Olfaction has been implicated in SHB infestations of honey bee and bumble bee colonies. We used olfactometer bioassays to investigate responses of adult male and female SHBs to odors from intact colonies and separate hive components (pot honey, pot pollen, cerumen, and propolis) of three African meliponine bee species, Meliponula ferruginea (Lepeletier) (black morphospecies), M. ferruginea (reddish brown morphospecies), and Meliponula bocandei (Spinola) (Hymenoptera: Apidae). Although both sexes of the beetle strongly preferred intact colony, pot honey, and pot pollen odors, there was no evidence of attraction to propolis and cerumen odors from the three meliponine bee species. Both sexes of SHB also strongly preferred odors from honey bees, Apis mellifera L. (Hymenoptera: Apidae), over odors from the three meliponine bee species. Our results provide substantial evidence of the host potential of African meliponine bees for the SHB, and we discuss this complex association of the SHB with species within the Apidae family.     

Honey bee (<Emphasis Type="Italic">Apis mellifera capensis/A. m. scutellata</Emphasis> hybrid) nesting behavior in the Eastern Cape,South Africa

Little is known about the natural history of wild honey bee (Apis mellifera) colonies in the Eastern Cape Province of South Africa. The goal of this research was to examine nest site characteristics of honey bee (A. m. capensis/A. m. scutellata hybrid) colonies sampled from a variety of habitats (nature reserves, livestock farms, and an urban setting) in the Eastern Cape. We also determined how nest site location related to various colony strength parameters. In general, colonies not nesting in ground cavities tended to nest in locations >6 m high when nesting in cliffs and buildings and >2 m high when nesting in trees. Colonies typically nested in cavities whose entrances faced a southeasterly direction and were ~40 L in volume. We sampled a subset of colonies to determine the relationship between nest type and the following colony strength parameters: total area of comb in the colony, the volume of stored honey, pollen, and brood, adult bee population, the weight per adult bee, and the bee/nest cavity volume ratio. In general, colonies nesting in cliffs tended to be stronger than those nesting in the ground or trees. Our findings provide new insights into the nesting biology of honey bees in the Eastern Cape, South Africa, perhaps leading to the formation of conservation recommendations for honey bees in this region.     

Western honey bee (Apis mellifera L.) attraction to commercial pollen substitutes and wildflower pollen in vitro

Many beekeepers feed their western honey bee (Apis mellifera) colonies artificial pollen substitutes to provide colonies with adequate nutrition during times of limited pollen quantity or quality. We provided caged worker bees commercially available pollen substitutes (AP23, MegaBee, UltraBee) and wildflower pollen in a choice-test to determine their relative attraction to/preference for the diets. We measured diet consumption by honey bees and observed honey bee behaviour to evaluate bee preferences for certain diets. The bees interacted with and consumed more wildflower pollen than they did any of the commercially available pollen substitutes. Our data suggest that bees have a strong preference for wildflower pollen over commercially available pollen substitutes.     

Microalgae as a promising and sustainable nutrition source for managed honey bees

Managed honey bee colony losses are attributed to a number of interacting stressors, but many lines of evidence point to malnutrition as a primary factor. Commercial beekeepers have become increasingly reliant on artificial pollen substitute diets to nourish colonies during periods of forage scarcity and to bolster colony size before pollination services. These artificial diets may be deficient in essential macronutrients (proteins, lipids, prebiotic fibers), micronutrients (vitamins, minerals), and antioxidants. Therefore, improving the efficacy of pollen substitutes can be considered vital to modern beekeeping. Microalgae are prolific sources of plant-based nutrition with many species exhibiting biochemical profiles that are comparable to natural pollen. This emerging feed source has been employed in a variety of organisms, including limited applications in honey bees. Herein, I introduce the nutritional value and functional properties of microalgae, extrapolating to central aspects of honey bee physiology and health. To conclude, I discuss the potential of microalgae-based feeds to sustainably provision managed colonies on an agricultural scale.     

The habitat disruption induces immune-suppression and oxidative stress in honey bees

The honey bee is a major insect used for pollination of many commercial crops worldwide. Although the use of honey bees for pollination can disrupt the habitat, the effects on their physiology have never been determined. Recently, honey bee colonies have often collapsed when introduced in greenhouses for pollination in Japan. Thus, suppressing colony collapses and maintaining the number of worker bees in the colonies is essential for successful long-term pollination in greenhouses and recycling of honey bee colonies. To understand the physiological states of honey bees used for long-term pollination in greenhouses, we characterized their gene expression profiles by microarray. We found that the greenhouse environment changes the gene expression profiles and induces immune-suppression and oxidative stress in honey bees. In fact, the increase of the number of Nosema microsporidia and protein carbonyl content was observed in honey bees during pollination in greenhouses. Thus, honey bee colonies are likely to collapse during pollination in greenhouses when heavily infested with pathogens. Degradation of honey bee habitat by changing the outside environment of the colony, during pollination services for example, imposes negative impacts on honey bees. Thus, worldwide use of honey bees for crop pollination in general could be one of reasons for the decline of managed honey bee colonies.     

Pollinator genetics and pollination: do honey bee colonies selected for pollen‐hoarding field better pollinators of cranberry Vaccinium macrocarpon?

1. Genetic polymorphisms of flowering plants can influence pollinator foraging but it is not known whether heritable foraging polymorphisms of pollinators influence their pollination efficacies. Honey bees Apis mellifera L. visit cranberry flowers for nectar but rarely for pollen when alternative preferred flowers grow nearby. 2. Cranberry flowers visited once by pollen‐foraging honey bees received four‐fold more stigmatic pollen than flowers visited by mere nectar‐foragers (excluding nectar thieves). Manual greenhouse pollinations with fixed numbers of pollen tetrads (0, 2, 4, 8, 16, 32) achieved maximal fruit set with just eight pollen tetrads. Pollen‐foraging honey bees yielded a calculated 63% more berries than equal numbers of non‐thieving nectar‐foragers, even though both classes of forager made stigmatic contact. 3. Colonies headed by queens of a pollen‐hoarding genotype fielded significantly more pollen‐foraging trips than standard commercial genotypes, as did hives fitted with permanently engaged pollen traps or colonies containing more larvae. Pollen‐hoarding colonies together brought back twice as many cranberry pollen loads as control colonies, which was marginally significant despite marked daily variation in the proportion of collected pollen that was cranberry. 4. Caloric supplementation of matched, paired colonies failed to enhance pollen foraging despite the meagre nectar yields of individual cranberry flowers. 5. Heritable behavioural polymorphisms of the honey bee, such as pollen‐hoarding, can enhance fruit and seed set by a floral host (e.g. cranberry), but only if more preferred pollen hosts are absent or rare. Otherwise, honey bees' broad polylecty, flight range, and daily idiosyncrasies in floral fidelity will obscure specific pollen‐foraging differences at a given floral host, even among paired colonies in a seemingly uniform agricultural setting.