A foraging insect has the potential to be a good pollinator, but not all foragers pollinate crops effectively. This could be due to foragers that are also flower eaters, foragers that are known as cheaters, or foragers that do not have the most efficient foraging physiology. The goal of using managed insects in a fruit crop, such as wild blueberries, is to find a foraging insect that will make the most efficient and effective pollination unit. Bees, such as honey bees, bumblebees, and alfalfa leafcutter bees, are the only managed pollinators that are used to pollinate wild blueberry crops in Atlantic Canada. Bees have many anatomical features that aid in foraging, and this is why bees are typically considered the most effective and efficient pollinators. 

Bee Foraging Physiology

The most efficient pollinator is the right shape and size, relative to the shape and size of the flowers they visit. This is because some parts of the pollinator’s body should be very close to or touching the anther of the flower to passively collect pollen during foraging activities [6]. The pollen that is actively collected is typically used by bees, so it is the passively collected pollen that gets transferred to the stigma of another flower for pollination. Flowers accumulate a negative charge and flying insects accumulate a positive charge, which allows these foragers to passively pick up pollen, through electroreception, as their body moves past anthers of a flower and carries it to another [1].

Pollen best adheres to fur or hair on a forager. If the surface of the forager is shiny, waxy, or contains chemicals then the pollen may not adhere to the forager or pollen could potentially be damaged, resulting in poor pollination [6]. Some bees, such as honey bees and bumblebees, can collect pollen on their corbiculae, which is a protrusion on the posterior of the hind tibia, also known as ‘pollen pants’. These bees collect pollen grains scattered on their bodies through different methods of grooming and pollen packing [8]. The methods used to groom and pack pollen by honey bees can damage pollen and reduce efficacy of pollination, unlike bumblebees. This is why passive pollen collection is important for successful pollination when using honey bees as pollinators. Bees may visit many flowers before returning to their hive, potentially pollinating flowers with the loose pollen on their body. 

Honey bee with 'pollen pants' on hind tibia (ATTTA ©, 2024)

Bees have two pairs of wings which are connected to work as a single pair of wings. These wings are attached by V-shaped hooks, called hamuli [5]. The hamuli are flexible, durable, and help increase surface area. These traits aid with flight, which helps the bees create additional lift when the bee is heavier with foraging resources and needs to travel long distances or avoid predators [9]. 

Honey bees are generalist foragers, meaning they collect pollen and nectar from many different floral sources. To be able to forage nectar with differing concentrations of sugar, bees can either suck or lap, depending on the viscosity of the nectar [11]. The fluid intake rate for sucking becomes more efficient than lapping at sugar concentrations lower than 30% [6]. Honey bees also have a crop, or honey stomach, to temporarily store honey to bring back to the hive, which can make up to a third of the bee’s weight [4,7]. Therefore, honey bees can travel from flower to flower, collecting a lot of nectar before going back home and this will also help with pollination efforts.

Honey bee foraging wild blueberry flower (ATTTA ©, 2024)

Insects of the Hymenoptera order, containing bee species, possess two to three times the number of genes for olfactory receptors than insects of the Diptera order, containing fly species [2]. Gene expression for olfactory receptors occurs in the bee’s information-gathering sense organ, the antenna [3]. Honey bees’ antennae are used to detect and respond to vibrations, electrical charges, and changes in temperature, humidity, and air quality, such as levels of carbon dioxide [1,6]. Bees also use the Organ of Johnston to detect sound and speed of movement, through changes in vibration [10]. This helps bees to adjust the pitch of their body and reduce drag while flying as well as detecting dances of other bees to find foraging resources.

Bees are typically considered the best pollinator due to the many anatomical traits they have to help with foraging. There are also many behavioral traits that bees have, to help with foraging, both on an individual and colony level. Please read the upcoming blog for more information on bee foraging behavior.

Written by John MacDonald, ATTTA Seasonal Apiculturist

Connecting with ATTTA Specialists

If you’d like to connect with ATTTA specialists or learn more about our program, you can:

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References: 

[1] Clarke, D., Morley, E., Robert, D. 2017. ‘The bee, the flower, and the electric field: electric ecology and aerial electroreception’, Journal of Comparative Physiology A, 203:737-748.

[2] De Bruyne, M., Baker, T.C., 2008. ‘Odor detection in insects: volatile codes’, Journal of Chemical Ecology, 34 (7):882–897.

[3] Fialho, M.Q., Guss-Matiello, C.P., Zanuncio, J.C., Campos, L.O., Serrão, J.E., 2015. ‘A comparative study of the antennal sensilla in corbiculate bees’, Journal of Apiculture Research, 53 (3):392–403.

[4] Harano, K.I., Mitsuhata-Asai, A., Konishi, T., Suzuki, T., Sasaki, M., 2013. ‘Honeybee foragers adjust crop contents before leaving the hive’, Behavioral Ecology and Sociobiology, 67 (7):1169–1178.

[5] Ma, Y., Ren, H., Ning, J., Gorb, S., 2022. ‘The combination of structure and material distribution ensures functionality of the honeybee wing-coupling mechanism’, Soft Matter 18 (5):956–963.

[6] Nearman, A., vanEngelsdorp, D. 2024. ‘Chapter 2 - Honey bee adaptations for foraging’, The Foraging Behavior of the Honey Bee (Apis mellifera)’, 45-64.

[7] Nicolson, S.W., Human, H., 2008. ‘Bees get a head start on honey production’, Biology Letters, 4:299–301.

[8] Parker, J.A., Tran, J.L., Ison, J.L., Bai, J.D., Weis, A.E., Thomson, J.D. 2015. ‘Pollen packing affects the function of pollen on corbiculate bees but not non-corbiculate bees’, Arthropod-Plant Interactions, 9:197-203. 

[9] Sudo, S., Tsuyuki, K., Ito, Y., Tani, J., Wada, H., 2001. ‘The wing apparatus and flapping behavior of Hymenoptera’, JSME International Journal Series C: Mechanical Systems, Machine Elements and Manufacturing, 44 (4):1103–1110.

[10] Taylor, G.J., Luu, T., Ball, D., Srinivasan, M.V., 2013. ‘Vision and air flow combine to streamline flying honeybees’, Scientific Reports, 3.

[11] Wei, J., Huo, Z., Gorb, S.N., Rico-Guevara, A., Wu, Z., Wu, J., 2020. ‘Sucking or lapping: facultative feeding mechanisms in honeybees (Apis mellifera)’, Biology Letters, 16 (8):20200449.

There are multiple forms of communication that honey bees utilize. Like many other animals, honey bees communicate with the use of vibrations and pheromones. But perhaps a more unique form of communication they use is dance. Honey bees have special dances performed by the worker bees that provide information about resources around the hive. Read this week’s blog to learn about the communicative dances honey bees perform.

Do the Waggle Dance 

There are a couple of special dances performed by worker bees to inform the hive about a food source and the location. The most famous of these dances is the waggle dance. Often when performing these dances, the workers provide a sample from the source to other foragers using trophallaxis. The returning forager also carries the smell of pollen and/or nectar, which provides more information about the food source to other bees.

The significance of the waggle dance is for foraging bees to notify other foragers when resources are more than 100 meters away from the hive. To notify, the bee moves in a figure eight pattern. More specifically, it is a straight run in the middle of the pattern with two semi-circle movements on either side. This provides information about the distance and direction of the resource (von Frisch, 1967). The waggle dance also informs bees of the quality of the resource (von Frisch, 1967).

There is also another dance that provides information about resources. The round dance informs bees when a resource is less than 15 meters away from a hive. The bee will dance in a round pattern, which tells other members of the colony that the resource is nearby. The round dance does not provide any information about the exact distance or direction of the resource (Frisch, 1967). However, that information is less essential when the resource is nearby.

There are some sceptics who believe that these dances may have no communication value, and that bees locate sources of food based on the scent left behind from previous foragers (Gadagkar, 1996). While bees can find food based on such odours, past experiments convincingly demonstrate that bees can also find food in the absence of smell. This once again suggests that bees find food based on information communicated through dance or possibly sound (Gadagkar, 1996). It has been suggested that honey bees recruit foragers using buzzing flights. These flights take place close to the resource and is another method bees utilize to locate food (Tautz and Sandeman, 2003).

Given that honey bees can find food sources by only using scent, it brings to question if dance communication provides an advantage to foraging bees. A study done by Okada et al. (2014) determined that colonies of honey bees that use the waggle dance have a significantly greater number of successful visits to food sources than colonies that do not use communication or use random dance communication. The experiment was performed with the use of virtual colonies and provides support that the waggle dance still has value in foraging ability.

Finally, several studies completed in the last decade suggest that human-modified habitats are having an impact on the value of dance language. Human-modified habitats are often characterized by mass-flowering crops that may be easy to find and profitable in spring.  However, once these crops have finished flowering, the environment becomes numerous low-quality foraging sites, and the dance’s value may be diminished (Nürnberger et al. 2017; Couvillon et al. 2014). These types of environments have many discrete foraging locations, but their quality is poor and the cost of recruitment to these sites outweighs the benefits. In summary, human impact may have created environments in which honey bee dance communication is not suited (Price, 2019).

Schematic of honey bee communicative dances (Frisch,1967)

Interestingly, honey bees are the only known group of bees that uses nest-based communication to provide information about food resources (Price and Grüter, 2015). The use of dance to communicate is just one of the reasons that honey bees are a highly productive and organized group.

Connecting with ATTTA Specialists

If you’d like to connect with ATTTA specialists or learn more about our program, you can:

visit our website at https://www.perennia.ca/portfolio-items/honey-bees/

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References
Couvillon, M.J., Fensome, K.A., Quah, S.K., and Schürch, R. 2014. Summertime blues: August foraging leaves honey bees empty-handed. Commun. Integr. Biol.
Frisch, Karl von. 1967. The Dance Language and Orientation of Bees. The Belknap Press of Harvard University Press, Cambridge, Massachusetts, pp. 566.
Gadagkar, R. 1996. The honeybee dance-language controversy. Resonance. 1(1): 63 – 70. doi:10.1007/bf02838860
Nürnberger, F., Steffan-Dewenter, I., and Härtel, S. 2017. Combined effects of waggle dance communication and landscape heterogeneity on nectar and pollen uptake in honey bee colonies. Peer J.
Price, R., Grüter, C. 2015. Why, when and where did honey bee dance communication evolve? Frontiers in Evolution and Ecology.
Price, R. et al. 2019. Honey bees forage more successfully without the “dance language” in challenging environments. Sci. Adv.
Tautz, J. and Sandeman, D.C. 2003. Recruitment of honeybees to non-scented food sources. J. Comp. Physiol. A Neuroethol. Sens. Neural. Behav. Physiol. 

In last week’s blog, we discussed the foraging behavior of honey bees while placed on almond orchards for pollination in Australia. In Atlantic Canada, honey bees are most heavily employed for the pollination of wild blueberries. Read on for insight into the foraging behavior of worker bees during this important period of the beekeeping season.

Honey Bee Nutrition in Wild Blueberry Fields

Providing wild blueberry pollination services is a major activity for many Atlantic Canadian beekeepers. Wild blueberries typically bloom in late May and early June, shortly after local forage sources have begun to proliferate. There are concerns that when bees are moved to blueberry fields, stress causes colonies to become weakened and susceptible to disease such as European foulbrood. In 2019, ATTTA did a study in Colchester county, Nova Scotia to explore this concern and improve our understanding of forage availability to honey bees (Olmstead et al. 2019).

Wild blueberry field in bloom. (ATTTA©2021)

ATTTA’s study explored nutritive health of pollinating honey bees by supplementally feeding and tracking hives during and after wild blueberry pollination. Seam counts and brood assessments revealed the colonies to be in good health. Colony growth and rate of EFB infection were not significantly different between hives fed pollen supplement and hives which were not. All colonies were at or above the Nova Scotia pollination standard for the duration of the trial and there were minimal instances of EFB.  In addition, pollen traps were mounted onto a sample of hives. After 24 hours, the collected pollen reflected a range of forage availability. 

The first published study to assess honey bee forage on wild blueberry fields in the Maritimes revealed that honey bees collected a very marginal amount of wild blueberry pollen during pollination (Colwell et al. 2017). This is distinct from honey bee pollination on almond orchards, where almond pollen is collected abundantly (Bezerra da Silva Santos et al. 2022). When honey bees are foraging on wild blueberry fields, they are nectar foraging (Javorek et al. 2002). They are moving from flower to flower, incidentally moving pollen in the process, but with the purpose of collecting nectar. Nectar is a valuable energy source for honey bees, while pollen provides protein and other nutrients that are important to honey bee health and brood rearing (Di Pasquale et al. 2013). When foraging wild blueberry fields, honey bees must rely on pollen from other resources.

ATTTA continued to sample pollen collections from honey bees during wild blueberry pollination in 2021 and 2022 to add to the breadth of knowledge that beekeepers and blueberry producers have about typical forage availability during this time. Stay tuned as we disseminate the results of these collections in upcoming meetings and publications. 

References

Bezerra da Silva Santos, Karen Cristine, Elizabeth Frost, Ulrika Samnegård, Manu E. Saunders, and Romina Rader. 2022. “Pollen Collection by Honey Bee Hives in Almond Orchards Indicate Diverse Diets.” Basic and Applied Ecology 64 (November): 68–78. https://doi.org/10.1016/j.baae.2022.07.006.
Colwell, Megan J., Geoffrey R. Williams, Rodger C. Evans, and Dave Shutler. 2017. “Honey Bee‐collected Pollen in Agro‐ecosystems Reveals Diet Diversity, Diet Quality, and Pesticide Exposure.” Ecology and Evolution 7 (18): 7243–53. https://doi.org/10.1002/ece3.3178.
Di Pasquale, Garance, Marion Salignon, Yves Le Conte, Luc P. Belzunces, Axel Decourtye, André Kretzschmar, Séverine Suchail, Jean-Luc Brunet, and Cédric Alaux. 2013. “Influence of Pollen Nutrition on Honey Bee Health: Do Pollen Quality and Diversity Matter?” Edited by Jochen Zeil. PLoS ONE 8 (8): e72016. https://doi.org/10.1371/journal.pone.0072016.
Javorek, S. K., K. E. Mackenzie, and S. P. Vander Kloet. 2002. “Comparative Pollination Effectiveness Among Bees (Hymenoptera: Apoidea) on Lowbush Blueberry (Ericaceae: <I>Vaccinium Angustifolium</I>).” Annals of the Entomological Society of America 95 (3): 345–51. https://doi.org/10.1603/0013-8746(2002)095[0345:CPEABH]2.0.CO;2.
Olmstead, Sawyer, Robyn McCallum, and Jillian Shaw. 2019. “Evaluating the Effect of Feeding Pollen Substitute to Honey Bee Colonies Destined for Wild Blueberry Pollinaiton in Cochester County, Nova Scotia.” Perennia.

Connecting with ATTTA Specialists

If you’d like to connect with ATTTA specialists or learn more about our program, you can:

visit our website at https://www.perennia.ca/portfolio-items/honey-bees/

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Honey bees are often used as either primary or supplemental pollinators in commercial agriculture. They are transported to flowering crop fields during the bloom period and subsequently returned to their home apiaries after providing pollination services. Honey bee nutrition depends on the colony’s surrounding floral sources, as this is where foragers can obtain pollen and nectar. Researchers in Australia have recently published a report which improves our knowledge of honey bee foraging behavior during crop pollination, read on to learn more about their encouraging findings. 

Honey Bee Nutrition in Commercial Pollination

In 2017, two Australian almond orchards were used as study sites to investigate the identity, quantity, and timing of pollen collection by honey bees used for pollination services (Bezerra da Silva Santos et al. 2022). In these orchards, beehives were distributed at several different locations within the fields. One orchard was stocked at 0.04 hives per hectare, holding 96 hives in 2,375 hectares, and the other was stocked at 0.06 hives per hectare, with 71 hives in a 1,200-hectare plot. Almond blossoms in this region are typically present in late August and early September for about two to four weeks. The colony placement preceded bloom by 7 days, and they then remained for 21 additional days of the full bloom period.  Pollen traps were placed on 80 colonies between the two fields and contents of trays were collected every 48 hours of the open bloom period. 

Foraging honey bees with yellow pollen packed in their corbiculae (ATTTA©2022).

This study demonstrated that honey bees collected an array of pollen varieties while placed on almond orchards. Seventy-nine percent of hives contained pollen from both non-almond and almond flowers.  This indicates that foragers were not limited to almond blossoms for nutrition. Almond pollen was most heavily collected at the beginning of the flowering period, with about 70% of all almond pollen collection being within the first three sampling days. This was the time at which there was the greatest number of open almond flowers. The researchers also found that the location within the orchards had no impact on the plant species of pollen collected. 

The results of this study are encouraging for beekeepers engaged in commercial pollination. It adds to the limited amount of research on honey bee nutrition during pollination. Wild blueberry pollination falls at a critical time for beekeepers in Atlantic Canada, as it is the beginning of our beekeeping season. There is nationwide concern among beekeepers regarding the health of their colonies during blueberry pollination, particularly related to nutritional stress (McAfee 2018). While wild blueberry pollination has important distinctions from almond pollination, the results of this study add to our general understanding of how honey bees forage on large, monoculture fields. This paper reflects that foragers are able to reach a variety of floral sources even in a vast orchard of almond blossoms. This is optimistic for beekeepers, as floral variety supports healthy honey bee nutrition (Donkersley et al. 2017).

In next week’s blog, we will discuss honey bee foraging as it relates more closely to wild blueberries and honey bee pollination in Atlantic Canada.   

References

Bezerra da Silva Santos, Karen Cristine, Elizabeth Frost, Ulrika Samnegård, Manu E. Saunders, and Romina Rader. 2022. “Pollen Collection by Honey Bee Hives in Almond Orchards Indicate Diverse Diets.” Basic and Applied Ecology 64 (November): 68–78. https://doi.org/10.1016/j.baae.2022.07.006.

Donkersley, Philip, Glenn Rhodes, Roger W. Pickup, Kevin C. Jones, Eileen F. Power, Geraldine A. Wright, and Kenneth Wilson. 2017. “Nutritional Composition of Honey Bee Food Stores Vary with Floral Composition.” Oecologia 185 (4): 749–61. https://doi.org/10.1007/s00442-017-3968-3.

McAfee, Ali. 2018. “The Blueberries and the Bees.” Bee Scene, 2018.

 Connecting with ATTTA Specialists

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visit our website at https://www.perennia.ca/portfolio-items/honey-bees/

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It is always lovely to see your garden admired by honeybees in the summer, buzzing around collecting the pollen and nectar. It is even nicer to see honeybees pollinating our orchards and field crops. Beyond the garden, managed honey bees are an agricultural necessity here in Atlantic Canada, pollinating everything from blueberry fields to apple orchards. However, many crops that are bee-pollinated only blossom for short periods and cannot sustain honey bees year-round. In this blog, we explore how beekeepers and non-beekeepers alike can support local honey bee populations by providing floral forage throughout the season.

Planting for the Bees

Nutritionally it is essential for bees to collect pollen and nectar from a variety of plants. It can be challenging for beekeepers to find suitable areas for hives that will fulfill both the type and quantity of blooming flowers that the bees need to stay healthy. Creating a garden design that will be in bloom from spring to fall is beneficial for foraging honey bees. The plants you choose to incorporate can be of any flowering variety, but consider native plants to provide a dependable and healthy ecosystem. There are many native trees and bushes such as maples, red-osier dogwood, ash trees, cherry, or oak. There are also many flowering plants, such as butterfly weed, common milkweed, bee borage, canola, lupine, fireweed, and so many more. For more ideas, visit the Atlantic Rhododendron and Horticultural Society website. This website includes a descriptive Excel table which lists over a hundred plants and when they bloom on the Atlantic coast. In addition to growing a variety of flowering plants, planting in clusters instead of spread-out patterns can make foraging more efficient for the bees. These flowering gardens will attract more than just honeybees. Also, consider adding old logs or rock piles as nesting features to support wild pollinators!

While it is valuable to plant new gardens for honey bee foraging, it is also good to consider the management of the wild flowers that are already present. Managed and wild growth is often controlled by mowing, which eliminates blooms, nesting resources, and pollinators' natural habitats. Depending on the time of mowing, the blades can also eliminate individual pollinators that may be present. Managing where and when you mow is an excellent course to ensure that foragers aren’t eliminated and get optimum use out of the plants which are present. Leaving patches unmowed and trimming as close to the roadside as possible can help. Leaving areas unmowed for as long as possible while plants are in bloom allows foragers to use up resources and then move onto new habitat.

Sustaining the health of our honeybees is growing more challenging as natural habitats are lost. Planting pollination gardens and managing native plants can help. The more flowers planted, the more food it provides, and the better the bees eat. When the bees are healthier, they are more equipped to fight off harmful pests and diseases, such as Varroa mites. This is why it is so important to consider planting for the bees. 

Written by Rebecca Campbell, ATTTA Summer Research Assistant rcampbell@perennia.ca

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This week, we will continue our discussion of wild blueberry pollination using honey bees. After stocking density, another consideration for wild blueberry producers is hive placement. It is important to have hives placed in an accessible location for beekeepers, and beyond that there are questions of spacing for maximum pollination. Today we will discuss hive placement on wild blueberry fields for optimal pollination services.

Hive Placement for Wild Blueberry Pollination

ATTTA began research in the 2021 field season to measure the distance that honey bees will successfully pollinate blueberry blooms in proximity to their hive. Our assessment included five points, the first of which was in the blueberry field near the recently placed hives.  From this point we continued across the field in a “W” shaped pattern to cover approximately +150m across the breadth of the field. At each of five field locations, we assessed 200 blueberry stems to establish the percent fruit set, percent of harvestable berries, and the berry weight over the course of three visits. 

In the ten fields we visited in both Nova Scotia and New Brunswick, we found no statistically significant difference in percent fruit set, harvestable berries, nor berry weight based on the distance of the sample location from the hives. This suggests that at a minimum of 150m from their hives, honey bees will evenly pollinate all blooms. Given these results, wild blueberry flowers should be consistently pollinated across a field if hives are placed in 300m increments.

Figure 1. Sampling sites (A – E) in the trial fields (n = 10) for three plant assessments. A) Fruit set as percent of bloom at all five field sampling sites. B) Fruit at harvest as a percent of blossom for all five field sampling sites. C) Final berry weight in grams just prior to harvest at all five There was no significant difference (p > 0.05) seen between any of the site locations across the three plant assessments.

Published research papers have also demonstrated that distance is not a limiting factor in successful honey bee pollination on wild blueberry fields, within the range of study. A simulation comparing the pollination services of honey bees, bumble bees, and solitary bees on fields in Maine demonstrated that unlike bumble bees and solitary bees, the distance of a honey bee hive from blueberry blooms did not significantly effect fruit set (Qu and Drummond 2018). In the simulation, bumble bee and solitary bee populations both showed reduced pollination activity, measured by fruit set, as the insects had to travel further from their hives and nests to pollinate clones. Honey bee foraging behavior is thought to play a role in this observation. When a foraging honey bee finds a source of nectar or pollen, she will return to the hive to tell other foragers about this source. In what is called the waggle dance, individual bees are able to provide other foragers with directions to this desirable location. As such, far away forage can be revisited multiple times and a honey bee colony can efficiently explore a large area (Tereshko and Loengarov 2005). 

Given that our results did not reach a limit of successful pollination, it is possible that honey bees will pollinate wild blueberry blooms beyond 150m from the hive. They can certainly be found at greater distances from their hives. An early study of honey bee pollination on wild blueberry fields in Quebec found that honey bee densities in a field did not begin to significantly drop until the hives were beyond 3km from the location in question (Aras, De Oliveira, and Savoie 1996). Before this point, honey bee density was measured to be well over 10 per 2m². 

The results of our research and relevant publications suggests that honey bees are steady pollinators in wild blueberry fields. Our results reflect that hive placement every 300 meters should provide consistent pollination throughout a field. Stay tuned for more as we carry on with our pollination series in a discussion of hive strength for pollination in a subsequent blog.

Aras, Philippe, Domingos De Oliveira, and Lorraine Savoie. 1996. “Effect of a Honey Bee (Hymenoptera: Apidae) Gradient on the Pollination and Yield of Lowbush Blueberry.” Journal of Economic Entomology 89 (5): 1080–83. https://doi.org/10.1093/jee/89.5.1080.

Qu, Hongchun, and Frank Drummond. 2018. “Simulation-Based Modeling of Wild Blueberry Pollination.” Computers and Electronics in Agriculture 144 (January): 94–101. https://doi.org/10.1016/j.compag.2017.11.003.

Tereshko, Valery, and Andreas Loengarov. 2005. “Collective Decision-Making in Honey Bee Foraging Dynamics,” 8.

Connecting with ATTTA Specialists

If you’d like to connect with ATTTA specialists or learn more about our program, you can:

visit our website at https://www.perennia.ca/portfolio-items/honey-bees/

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