other workers to help develop queen cups during the process of queen rearing.
A honey bee colony has a social organization characterized by a division of labor among the worker honey bees – worker polyethism – that changes over time (Winston 1987; Seeley 1995). During their early life (0–20 days) they work inside the colony cleaning cells and caring for the brood, receiving nectar and handling pollen, comb building, and tending the queen among other tasks; as they age (20–45 days) they begin working outside the nest ventilating the hive and guarding its entrance, as well as commencing foraging flights (Winston 1987). The control of worker polyethism has been a mystery, yet recent evidence suggests that a pheromone, produced by older foraging sister worker bees, delays the age of onset of foraging and other age‐related tasks (Leoncini et al. 2004; Slessor et al. 2005).
Drone Pheromones
The male or drone honey bee differs significantly from his worker sisters, and with their singular reproductive role of sperm production and mating, drones produce very few pheromonal signals (Bortolotti and Costa 2014; Hrassnigg and Crailsheim 2005). The most important pheromone of drone bees is released from their mandibular gland, and functions to attract other drone bees to the male congregation areas in preparation for mating. Similar to female honey bees, drone bees also possess unique cuticular pheromone signals that allow worker bees to distinguish between the sexes and drone bees of different ages (Bortolotti and Costa 2014). In addition, the regulation of drone brood production and the rejection of adult drone bees from a colony are likely under pheromonal control (Free 1987).
Brood Pheromones
Larvae within the honey bee colony release pheromones that are important for the regulation of brood care and development, worker behavioral transformation, and worker reproduction (Bortolotti and Costa 2014; Free 1987). Different components of the brood pheromone are released as a function of the caste and larval age, and as such guide the nurse bees to provide the appropriate response during brood development (Bortolotti and Costa 2014). Brood pheromone affects the colony foraging behavior in a dose‐dependent manner according to the age of the larvae. Young larvae (having little nursing needs) stimulate foraging and pollen collection, while older larvae (having greater nursing needs) delay foraging and instead promote increased brood care (Bortolotti and Costa 2014). Honey bee eggs, larvae, and pupae also stimulate pollen collection, and such brood pheromones are important modulators of colony growth (Bortolotti and Costa 2014; Free 1987). Further, brood pheromones work together with the queen signal to inhibit worker ovary development.
Acoustic Communication
Most of the social life of honey bees occurs within the darkness of the hive where vision plays a limited role (Kirchner 1993). Undeniably, pheromone communication provides the foundation for communication throughout this dark world of the honey bee (Slessor et al. 2005). However, honey bees can also detect and communicate via sound and vibrations that are transmitted throughout the beeswax structure of the hive (Hrncir et al. 2005; Kirchner 1993). Several types of acoustic communications are known within the honey bee colony and such signals may be transmitted via both substrate vibrations and airborne sound. These include the “tooting” and “quacking” signals among queens during the course of swarming, worker piping through dance language and swarming, as well as the hissing or shimmering behavior of honey bees during colony defense (Hrncir et al. 2005). Even though our knowledge of the complex interactions of the chemical, tactile, and acoustic sounds exploited by honey bees continues to advance, parts of their fascinating social lifestyle remain yet unknown.
Conclusion
When beekeepers and veterinarians understand the fascinating biology of the honey bee, including their anatomy and physiology, they will learn to appreciate the importance of the individual bee and the collective colony. They will both share a respect for the intricate and complex work, actions, and behaviors that allow thousands of relatives to share a common home and protect a cooperative future. And perhaps most importantly, the beekeeper and their veterinary colleague will build a foundation for effective communication, intervention and prevention of management problems and disease, helping to secure the future of this essential resource.
References
1 Amdam, G.V., Fennern, E., and Havukainen, H. (2012). Vitellogenin in honey bee behavior and lifespan. In: Honeybee Neurobiology and Behavior: A Tribute to Randolf Menzel (eds. C.G. Galizia, D. Eisenhardt and M. Giurfa). Switzerland: Springer Nature.
2 Arrese, E.L. and Soulages, J.L. (2010). Insect fat body: energy, metabolism, and regulation. Annual Review of Entomology 55: 207–225.
3 Bell, W.J. and Carde &c.acute;, R.T. (1984). Preface, Page xiv. In: Chemical Ecology of Insects (eds. W.J. Bell and R.T. Carde &c.acute;). London: Chapman and Hall.
4 Bortolotti, L. and Costa, C. (2014). Chemical communication in the honey bee society Chapter 5. In: Neurobiology of Chemical Communication (ed. C. Mucignat‐Caretta). Boca Raton, FL: CRC Press/Taylor & Francis.
5 Breed, M.D., Guzman‐Novoa, E., and Hunt, G.J. (2004). Defensive behavior of honey bees: organization, genetics and comparisons with other bees. Annual Review of Entomology 49: 271–298.
6 Brockmann, A. and Robinson, G.E. (2007). Central projections of sensory systems involved in honey bee dance language communication. Brain Behav Evol 70: 125–136.
7 Corona, M., Velarde, R.A., Remolina, S. et al. (2007). Vitellogenin, juvenile hormone, insulin signaling and queen bee longevity. Proceedings of the National Academy of Sciences of the United States of America 104 (17): 7128–7133.
8 Davidson, E.W. (1973). Ultrastructure of American foulbrood disease pathogenesis in larvae of the worker honey bee, Apis mellifera. Journal of Invertebrate Pathology 21: 53–61.
9 DeGrandi‐Hoffman, G. and Chen, Y. (2015). Nutrition, immunity and viral infections in honey bees. Current Opinion in Insect Science 10: 170–176.
10 Döke, M.A., Frazier, M., and Grozinger, C.M. (2015). Overwintering honey bees: biology and management. Current Opinion in Insect Science 10: 185–193.
11 Dor, R., Katzav‐Gozansky, T., and Hefetz, A. (2005). Dufour's gland pheromone as a reliable fertility signal among honeybee (Apis mellifera) workers. Behavioral Ecology and Sociobiology 58 (3): 270–276.
12 Elekonich, M.M. and Roberts, S.P. (2005). Honey bees as a model for understanding mechanisms of life history transitions. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 141: 362–371.
13 Ellis, J.D. (2015). The internal anatomy of the honey bee. American Bee Journal: 971–974.
14 Evans, J.D., Aronstein, K., Chen, Y.P. et al. (2006). Immune pathways and defense mechanisms in honey bees Apis mellifera. Insect Molecular Biology 15 (5): 645–656.
15 Fefferman, N.H., Traniello, J.F.A., Rosengaus, R.B., and Calleri, D.V. (2007). Disease prevention and resistance in social insects: modeling the survival consequence of immunity, hygienic behavior and colony organization. Behavioral Ecology and Sociobiology 61 (4): 565–577.
16 Fluri, P., Lüscher, M., Wille, H., and Gerig, L. (1982). Changes in weight of the pharyngeal gland and haemolymph titers of juvenile hormone, protein and vitellogenin in worker honey bees. Journal of Insect Physiology 28 (1): 61–68.
17 Free, J.B. (1987). Pheromones of Social Bees. Ithaca, NY: Cornell University Press.
