, Volume 101, Issue 10, pp 783–790 | Cite as

A critical number of workers in a honeybee colony triggers investment in reproduction

  • Michael L. Smith
  • Madeleine M. Ostwald
  • J. Carter Loftus
  • Thomas D. Seeley
Original Paper


Social insect colonies, like individual organisms, must decide as they develop how to allocate optimally their resources among survival, growth, and reproduction. Only when colonies reach a certain state do they switch from investing purely in survival and growth to investing also in reproduction. But how do worker bees within a colony detect that their colony has reached the state where it is adaptive to begin investing in reproduction? Previous work has shown that larger honeybee colonies invest more in reproduction (i.e., the production of drones and queens), however, the term ‘larger’ encompasses multiple colony parameters including number of adult workers, size of the nest, amount of brood, and size of the honey stores. These colony parameters were independently increased in this study to test which one(s) would increase a colony’s investment in reproduction via males. This was assayed by measuring the construction of drone comb, the special type of comb in which drones are reared. Only an increase in the number of workers stimulated construction of drone comb. Colonies with over 4,000 workers began building drone comb, independent of the other colony parameters. These results show that attaining a critical number of workers is the key parameter for honeybee colonies to start to shift resources towards reproduction. These findings are relevant to other social systems in which a group’s members must adjust their behavior as a function of the group’s size.


Sociogenesis Reproductive investment Reproductive timing Worker number Honeybees Drone comb 



We thank Kevin Loope, Julie Miller, and Carmen Kelleher, for their critical readings of the manuscript, and Jason Barry (CSCU), for his statistical advice. This paper is based on work supported by a US National Science Foundation Graduate Research Fellowship (to MLS) and by a Hatch Grant (2010-11-237) from the Cornell University Agriculture Experiment Station (to TDS).


  1. Akaike H (1974) A new look at the statistical model identification. Autom Control IEEE Trans 19:716–723. doi: 10.1109/TAC.1974.1100705 CrossRefGoogle Scholar
  2. Alaux C, Jaisson P, Hefetz A (2005) Reproductive decision-making in semelparous colonies of the bumblebee Bombus terrestris. Behav Ecol Sociobiol 59:270–277. doi: 10.1007/s00265-005-0035-6 CrossRefGoogle Scholar
  3. Boes KE (2010) Honeybee colony drone production and maintenance in accordance with environmental factors: an interplay of queen and worker decisions. Insect Soc 57:1–9. doi: 10.1007/s00040-009-0046-9 CrossRefGoogle Scholar
  4. Brian MV (1957) The growth and development of colonies of the ant myrmica. Insect Soc 4:177–190CrossRefGoogle Scholar
  5. Chien AC, Hill NS, Levin PA (2012) Cell size control in bacteria. Curr Biol 22:R340–R349. doi: 10.1016/j.cub.2012.02.032 PubMedCentralPubMedCrossRefGoogle Scholar
  6. Cole BJ, Wiernasz DC (2000) Colony size and reproduction in the western harvester ant, Pogonomyrmex occidentalis. Insect Soc 47:249–255. doi: 10.1007/PL00001711 CrossRefGoogle Scholar
  7. Free J, Williams I (1975) Factors determining the rearing and rejection of drones by the honeybee colony. Anim Behav 23:650–675CrossRefGoogle Scholar
  8. Gordon DM, Paul RE, Thorpe K (1993) What is the function of encounter patterns in ant colonies? Anim Behav 45:1083–1100. doi: 10.1006/anbe.1993.1134 CrossRefGoogle Scholar
  9. Grozinger CM, Richards J, Mattila HR (2013) From molecules to societies: mechanisms regulating swarming behavior in honey bees (Apis spp.). Apidologie 45:327–346. doi: 10.1007/s13592-013-0253-2 CrossRefGoogle Scholar
  10. Henderson CE (1991) Reproductive investment in drones in honey bee (Apis mellifera) colonies. Dissertation, Cornell UniversityGoogle Scholar
  11. Imdorf A, Buehlmann G, Gerig L et al (1987) Überprüfung der Schätzmethode zur Ermittlung der Brutfläche und der Anzahl Arbeiterinnen in freifliegenden Bienenvölkern. Apidologie 18:137–146CrossRefGoogle Scholar
  12. Jeanne RL, Bouwma AM (2002) Scaling in nests of a social wasp: a property of the social group. Biol Bull 202:289–295PubMedCrossRefGoogle Scholar
  13. Kozlowski J (1992) Optimal allocation of resources to growth and reproduction: implications for age and size at maturity. Trends Ecol Evol 7:15–19. doi: 10.1016/0169-5347(92)90192-E PubMedCrossRefGoogle Scholar
  14. Lee P, Winston ML (1985) The effect of swarm size and date of issue on comb construction in newly founded colonies of honeybees (Apis mellifera L.). Can J Zool 63:524–527CrossRefGoogle Scholar
  15. Martin H, Lindauer M (1966) Sinnesphysiologische Leistungen beim Wabenbau der Honigbiene. Z Vergl Physiol 53:372–404CrossRefGoogle Scholar
  16. Michener CD (1964) Reproductive efficiency in relation to colony size in hymenopterous societies. Insect Soc 11:317–342CrossRefGoogle Scholar
  17. Mitchell C (1970) Weights of workers and drones. Am Bee J 110:468–469Google Scholar
  18. Muller CB, Schmid-Hempel P (1992) Variation in life-history pattern in relation to worker mortality in the bumble-bee, Bombus lucorum. Funct Ecol 6:48–56CrossRefGoogle Scholar
  19. Otis GW (1982) Weights of worker honeybees in swarms. J Apic Res 21:88–92Google Scholar
  20. Page RE (1981) Protandrous reproduction in honey bees. Environ Entomol 10:359–362Google Scholar
  21. Pinter-Wollman N, Bala A, Merrell A et al (2013) Harvester ants use interactions to regulate forager activation and availability. Anim Behav 86:197–207. doi: 10.1016/j.anbehav.2013.05.012 PubMedCentralPubMedCrossRefGoogle Scholar
  22. Pomeroy N, Plowright R (1982) The relation between worker numbers and the production of males and queens in the bumble bee Bombus perplexus. Can J Zool 60:954–957CrossRefGoogle Scholar
  23. Pratt SC (1999) Optimal timing of comb construction by honeybee (Apis mellifera) colonies: a dynamic programming model and experimental tests. Behav Ecol Sociobiol 46:30–42. doi: 10.1007/s002650050589 CrossRefGoogle Scholar
  24. R Core Team (2012) R: A language and environment for statistical computing.
  25. Rangel J, Seeley TD (2012) Colony fissioning in honey bees: size and significance of the swarm fraction. Insect Soc 59:453–462. doi: 10.1007/s00040-012-0239-5 CrossRefGoogle Scholar
  26. Seeley TD (1989) The honey bee colony as a superorganism. Am Sci 77:546–553Google Scholar
  27. Seeley TD, Tautz J (2001) Worker piping in honey bee swarms and its role in preparing for liftoff. J Comp Physiol A 187:667–676. doi: 10.1007/s00359-001-0243-0 PubMedCrossRefGoogle Scholar
  28. Starr CK (2006) Steps toward a general theory of the colony cycle in social insects. In: Kipyatkov VE (ed) Life cycles in social insects: behaviour, ecology and evolution. St. Petersburg University Press, St. Petersburg, pp 1–20Google Scholar
  29. Taber S, Owens CD (1970) Colony founding and initial nest design of honey bees, Apis mellifera L. Anim Behav 18:625–632CrossRefGoogle Scholar
  30. Thummel CS (2001) Molecular mechanisms of developmental timing in C. elegans and Drosophila. Dev Cell 1:453–465PubMedCrossRefGoogle Scholar
  31. Tschinkel WR (1991) Insect sociometry, a field in search of data. Insect Soc 38:77–82. doi: 10.1007/BF01242715 CrossRefGoogle Scholar
  32. Tschinkel WR (1993) Sociometry and sociogenesis of colonies of the fire ant Solenopsis invicta during one annual cycle. Ecol Soc Am 63:425–457Google Scholar
  33. Tschinkel WR (1998) Sociometry and sociogenesis of colonies of the harvester ant, Pogonomyrmex badius: worker characteristics in relation to colony size and season. Insect Soc 45:385–410. doi: 10.1007/s000400050097 CrossRefGoogle Scholar
  34. Tschinkel WR (2011) Back to basics: sociometry and sociogenesis of ant societies (Hymenoptera: Formicidae). Myrmecol News 14:49–54Google Scholar
  35. Turner JJ, Ewald JC, Skotheim JM (2012) Cell size control in yeast. Curr Biol 22:R350–R359. doi: 10.1016/j.cub.2012.02.041 PubMedCentralPubMedCrossRefGoogle Scholar
  36. Webb M (1961) The biology of the bumblebees of a limited area in eastern Nebraska. University of Nebraska, DissertationGoogle Scholar
  37. Wilson EO (1985) The sociogenesis of insect colonies. Science 228:1489–1495PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  • Michael L. Smith
    • 1
  • Madeleine M. Ostwald
    • 1
  • J. Carter Loftus
    • 1
  • Thomas D. Seeley
    • 1
  1. 1.Department of Neurobiology and BehaviorCornell UniversityIthacaUSA

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