Honeybee queens

The repertoire of chemicals that can be used for communication is limited by the biosynthetic ability of the insect. Compared to other insect orders, pheromone biosynthesis in Hymenoptera has received little study [191]. However, the biosynthetic origins of chemically diverse hymenopteran semiochemicals likely include aromatic, fatty acid, and terpenoid pathways as well as simple modifications of host-derived precursors. Notable recent studies include the biosynthesis of the fatty acid components (2 )-9-oxodec-2-enoic acid 52 and (2 )-9-hydroxydec-2-enoic acid of the honeybee queen mandibular pheromone from octadecanoic acid [192,193], and the aliphatic alcohol and ester... 

Slessor KN,Foster LJ,Winston ML (1998) Royal flavours honeybee queen pheromones. In Vander Meer RK, Breed MD, Espelie KE, Winston ML (eds) Pheromone communication in social insects ants, wasps, bees and termites. Westview Press, Boulder, Colorado 331... 

A similar approach was applied in the study of the silkmoth bombykol pheromone receptor (Sakurai et al. 2004 Nakagawa et al. 2005), a receptor for honeybee queen substance p-oxo-2-decenoic acid (Wanner et al. 2007), and a ligand repertoire for Anopheles ORs (Lu et al. 2007). In these cases, however, coexpression of Ga15 was not necessary, but expression of one member of the Or83b family was absolutely necessary. The molecular mechanisms for signal transduction are discussed in Sect. 4. 

Petroleum is not the only place where alkanes occur naturally. Solid n-alkanes, especially those with relatively long chains, have a waxy constituency and coat the outer surface of many living things where they help prevent the loss of water. Pentacosane [CH3(CH2)23CH3] is present in the waxy outer layer of most insects. Hentriacontane [CH3(CH2)29CH3] is a component of beeswax (see Problem 2.6) as well as the wax that coats the leaves of tobacco, peach trees, pea plants, and numerous others. The C23, C25, C27, C29, and C31 /2-alkanes have been identified in the surface coating of the eggs of honeybee queens. 

Figure 4, Power-time curve of a virgin honeybee queen Apis mellifera camica) at 25 °C together with 3 frequency spectra for the points a, b and c. Besides the ground noise, there are nearly no higher frequencies for point a of resting metabolism, while at b and c higher frequencies dominate prior to the heat increase in the curve [57],...

Among the social Hymenoptera, one of the best-understood examples is the sex pheromone used by the queen honeybee to attract drones for mating. This pheromone is produced in the mandibular gland that resides in the head capsule (Barbier and Lederer, 1960 Callow and Johnson, 1960 Callow et al., 1964). In... 

Callow R. K. and Johnson N. C. (1960) The chemical constitution and synthesis of queen substance of honeybees (Apis mellifera L.). Bee World 41, 152-153. 

The best understood of the sexual pheromones of social insects is the queen substance of honeybees. Interestingly, the queen substance used for queen control inside the nest is also the substance used by virgin queens to attract drones for mating. Callow and Johnston (1960) and Barbier and Lederer (1960) identified 9-keto-2(E)-decenoic acid ([E]-9-oxodec-2-enoic acid) (9-ODA) as major components of the queen mandibular glands. 9-Hydroxy-2(E)-decenoic acid is also present (Callow et al., 1964) and together both attract drones. Additional components of the queen retinue pheromone have recently been identified (Keeling et al., 2003). 

Figure 11.2 Biosynthetic pathways leading to honeybee worker and queen mandibular gland pheromones. Reprinted with permission from Plettner E., et al. (1966), 1966 American Association for the Advancement of Science.

Katzav-Gozansky T., Soroker V., Ibarra F., Francke W. and Hefetz A. (2001a) Dufour s gland secretion of the queen honeybee (Apis mellifera) an egg discriminator or a queen signal Behav. Ecol. Sociobiol. 51, 76-86. 

Moritz R. F. A., Crewe R. M. and Hepburn H. R. (2002) Queen avoidance and mandibular gland secretion of honeybee workers (Apis mellifera L.). Insectes Soc. 49 86-91. 

Pankiw T., Winston M. L., Fondrk M. K. and Slessor K. N. (2000) Selection on worker honeybee responses to queen pheromone (Apis mellifera L.). Naturwissenschaften 87, 487 190. 

Danty E., Briand L., Michard-Vanhee C., Perez V., Arnold G. and Gaudemer O., Huet D., Huet J. C., Ouali C., Masson C. and Pernollet J. C. (1999) Cloning and expression of a queen pheromone-binding protein in the honeybee an olfactory-specific, developmentally regulated protein. J. Neurosci. 19, 7468-7475. 

Despite the importance of such pheromones for the understanding of insect sociality, very little is yet known about them (Le Conte and Hefetz, 2008). So far, in only one species (i.e. the honeybee), has it been shown that a pheromone emitted by a reproductive prevents worker reproduction. In the honeybee, Apis mellifera, it has been suggested that the queen mandibular pheromone (QMP) causes worker ovarian inhibition (Butler and Fairey, 1963). 

It is also important to consider other potential sources that may contribute to the recognition of reproductive individuals. So far, the vast majority of studies indicate a major role for long-chain hydrocarbons, but polar compounds may occasionally be involved as well (Tentschert et al., 2002 Sramkova et al., 2008). These polar compounds may also be of proteinaceous nature. They are found to differ between foundresses in Polistes dominulus (Dapporto et al., 2008). Although polar compounds are involved in the inhibition of ovarian activity in honeybee workers (Hoover et al., 2003) and prevent intracolonial reproduction of new queens in the ant Solenopsis invicta (Vargo, 1997,1998), these findings seem to be exceptions. 

Katzav-Gozansky, T., Soroker, V., Kamer, J., Schulz, C.M., Francke, W. and Hefetz, A. (2003). Ultrastructural and chemical characterization of egg surface of honeybee worker and queen-laid eggs. Chemoecology, 13, 129-134. 

Dietemann, V., Pflugfelder, J., Hartel, S., Neumann, P. and Crewe, R.M. (2006). Social parasitism by honeybee workers (Apis mellifera capensis Esch.) evidence for pheromonal resistance to host queen s signals. Behav. Ecol. Sociobiol., 60, 785-793. 

Primer pheromones, which cause long-term behavioral changes, are harder to isolate and identify. One example, however, is the queen substance produced by queen honeybees. All the eggs in a colony are laid by one queen bee. If she is removed from the hive or dies, the worker bees are activated by the absence of the queen substance and begin to feed royal jelly to bee larvae to raise a new queen. The queen substance also prevents the development... 

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