Behavioral Ecology and Sociobiology

, Volume 65, Issue 2, pp 305–312 | Cite as

Deciding when to explore and when to persist: a comparison of honeybees and bumblebees in their response to downshifts in reward

  • John M. Townsend-MehlerEmail author
  • Fred C. Dyer
  • Kim Maida
Original Paper


As the distribution of food resources shifts over time, central place foragers are likely to be repeatedly faced with the question of when to abandon a forage site that is declining in value and to subsequently search elsewhere. Although there has been a great deal of investigation into how individual foragers allocate time between exploration and exploitation, few studies have sought to explore this issue within a larger functional context. We take a comparative approach to this problem by examining decision making in individual honeybees and bumblebees as they responded to a downshift in food reward. Our results show not only that honeybees and bumblebees have significantly divergent strategies with regards to abandoning a food source that is declining in value but also in terms of the subsequent tendency to seek an alternative food source. We interpret these results in terms of both biological and social distinctions between these species and highlight how group-level characteristics are likely to shape the evolutionarily derived foraging strategies of individual animals.


Foraging behavior Honeybees Bumblebees Exploration versus exploitation Social foraging 



We would like to thank Mara Trudgen and Michael Hillman for their all help with data collection. We would also especially like to thank Martin Giurfa and two anonymous reviewers for their enormously helpful comments on the manuscript.


  1. Azoulay-Schwartz R, Kraus S, Wilkenfeld J (2004) Exploitation vs. exploration: choosing a supplier in an environment of incomplete information. Decis Support Syst 38:1–18CrossRefGoogle Scholar
  2. Blumstein DT, Daniel JC (2007) Quantifying behavior the JWatcher way. Sinauer Associates, SunderlandGoogle Scholar
  3. Cartar RV, Abrahams MV (1996) Risk-sensitive foraging in a patch departure context: a test with worker bumble bees. Am Zool 36:447–458Google Scholar
  4. Charnov EL (1976) Optimal foraging, the marginal value theorem. Theor Popul Biol 9:129–136CrossRefPubMedGoogle Scholar
  5. Clements KM, Wainwright PE (2007) Spontaneously hypertensive, Wistar Kyoto and Sprague-Dawley rats differ in performance on a win-stay task and a conditioned cue preference task in the water radial arm maze. Behav Brain Res 183:169–177CrossRefPubMedGoogle Scholar
  6. Cnaani J, Schmid-Hempel R, Schmidt JO (2002) Colony development, larval development and worker reproduction in Bombus impatiens Cresson. Insect Soc 49:164–170CrossRefGoogle Scholar
  7. Cohen JD, McClure SM, Yu AJ (2007) Should I stay or should I go? How the human brain manages the trade-off between exploitation and exploration. Philos Trans R Soc B 362:933–942Google Scholar
  8. Daw ND, O'Doherty JP, Dayan P, Seymour B, Dolan RJ (2006) Cortical substrates for exploratory decisions in humans. Nature 441:876–879CrossRefPubMedGoogle Scholar
  9. Dornhaus A, Chittka L (2001) Food alert in bumblebees (Bombus terrestris): possible mechanisms and evolutionary implications. Behav Ecol Sociobiol 50:570–576CrossRefGoogle Scholar
  10. Dornhaus A, Chittka L (2004) Why do honey bees dance? Behav Ecol Sociobiol 55:395–401CrossRefGoogle Scholar
  11. Driessen G, Bernstein C (1999) Patch departure mechanisms and optimal host exploitation in an insect parasitoid. J Anim Ecol 68:445–459CrossRefGoogle Scholar
  12. Dyer FC (2002) The biology of the dance language. Annu Rev Entomol 47:917–949CrossRefPubMedGoogle Scholar
  13. Dyer AG, Spaethe J, Prack S (2008) Comparative psychophysics of bumblebee and honeybee colour discrimination and object detection. J Comp Physiol Neuroethol Sens Neural Behav Physiol 194:617–627CrossRefGoogle Scholar
  14. Frisch KV (1993) The dance language and orientation of bees. Harvard University Press, CambridgeGoogle Scholar
  15. Giurfa M (1996) Movement patterns of honeybee foragers: motivation and decision rules dependent on the rate of reward. Behaviour 133:579–596CrossRefGoogle Scholar
  16. Goulson D, Peat J, Stout JC, Tucker J, Darvill B, Derwent LC, Hughes WOH (2002) Can alloethism in workers of the bumblebee, Bombus terrestris, be explained in terms of foraging efficiency? Anim Behav 64:123–130CrossRefGoogle Scholar
  17. Greggers U, Menzel R (1993) Memory dynamics and foraging strategies of honeybees. Behav Ecol Sociobiol 32:17–29CrossRefGoogle Scholar
  18. Heinrich B (1979a) Bumblebee economics. Harvard University Press, CambridgeGoogle Scholar
  19. Heinrich B (1979b) Majoring and minoring by foraing bumblebees, Bombus vegans—experimental analysis. Ecology 60:245–255CrossRefGoogle Scholar
  20. Heinrich B (2004) Bumblebee economics. Harvard University Press, CambridgeGoogle Scholar
  21. Hodges CM (1985) Bumble bee foraging—the threshold departure rule. Ecology 66:179–187CrossRefGoogle Scholar
  22. Kaelbling LP, Littman ML, Moore AW (1996) Reinforcement learning: a survey. J Artif Intell Res 4:237–285Google Scholar
  23. Krebs JR, Ryan JC, Charnov EL (1974) Hunting by expectation or optimal foraging—study of patch use by chickadees. Anim Behav 22:953CrossRefGoogle Scholar
  24. Macuda T, Gegear RJ, Laverty TM, Timney B (2001) Behavioural assessment of visual acuity in bumblebees (Bombus impatiens). J Exp Biol 204:559–564PubMedGoogle Scholar
  25. Maeda T, Takabayashi J (2001) Patch-leaving decision of the predatory mite Amblyseius womersleyi (Acari: Phytoseiidae) based on multiple signals from both inside and outside a prey patch. J Insect Behav 14:829–839CrossRefGoogle Scholar
  26. Matsen FA, Nowak MA (2004) Win-stay, lose-shift in language learning from peers. Proc Natl Acad Sci USA 101:18053–18057CrossRefPubMedGoogle Scholar
  27. McNamara J (1982) Optimal patch use in a stocastic environment. Theor Popul Biol 21:269–288CrossRefGoogle Scholar
  28. Michener CD (2000) The bees of the world. Johns Hopkins University Press, BaltimoreGoogle Scholar
  29. Nicolson SW, Nepi M, Pacini E (2007) Nectaries and nectar. Springer, DordrechtCrossRefGoogle Scholar
  30. Pyke GH (1978) Optimal foraging—movement patterns of bumblebees between inflorescences. Theor Popul Biol 13:72–98CrossRefPubMedGoogle Scholar
  31. Schaffer WM, Jensen DB, Hobbs DE, Gurevitch J, Todd JR, Schaffer MV (1979) Competition, foraging energetics, and the cost of sociality in 3 speices of bees. Ecology 60:976–987CrossRefGoogle Scholar
  32. Scheiner R, Page RE, Erber J (2004) Sucrose responsiveness and behavioral plasticity in honey bees (Apis mellifera). Apidologie 35:133–142CrossRefGoogle Scholar
  33. Schmid-Hempel P, Kacelnik A, Houston AI (1985) Honeybeees maximize efficiency by not filling their crop. Experientia 41:1229Google Scholar
  34. Seeley TD (1986) Social foraging by honeybees—how colonies allocate foragers among patches of flowers. Behav Ecol Sociobiol 19:343–354CrossRefGoogle Scholar
  35. Seeley TD (1989) Social foraging in honey bees—how nectar foragers assess their colony nutritional status. Behav Ecol Sociobiol 24:181–199CrossRefGoogle Scholar
  36. Seeley TD (1994) Honey-bee foragers as sensory units of their colonies. Behav Ecol Sociobiol 34:51–62CrossRefGoogle Scholar
  37. Seeley TD (1995) The wisdom of the hive: the social physiology of honey bee colonies. Harvard University Press, CambridgeGoogle Scholar
  38. Seeley TD, Camazine S, Sneyd J (1991) Collective decision-making in honey-bees—how colonies choose among nectar sources. Behav Ecol Sociobiol 28:277–290CrossRefGoogle Scholar
  39. Shimp CP (1966) Probabilistically reinforced choice behavior in pigeons. J Exp Anal Behav 9:443CrossRefPubMedGoogle Scholar
  40. Suarez RK, Lighton JRB, Joos B, Roberts SP, Harrison JF (1996) Energy metabolism, enzymatic flux capacities, and metabolic flux rates in flying honeybees. Proc Natl Acad Sci USA 93:12616–12620CrossRefPubMedGoogle Scholar
  41. Towne WF, Gould JL (1988) The spatial precisions of the honey bees’ dance communication. J Insect Behav 1:129–155CrossRefGoogle Scholar
  42. Townsend-Mehler JM (2010) Decision-making in a changing environment: a look at the foraging behavior of honeybees and bumblebees as they respond to shifts in resource availability. PhD thesis. Michigan State UniversityGoogle Scholar
  43. Waage JK (1979) Foraging for patchily-distributed hosts by the parasitoid, Nemeritis canescens. J Anim Ecol 48:353–371CrossRefGoogle Scholar
  44. Wajnberg E, Bernhard P, Hamelin F, Boivin G (2006) Optimal patch time allocation for time-limited foragers. Behav Ecol Sociobiol 60:1–10CrossRefGoogle Scholar
  45. Warrant E, Porombka T, Kirchner WH (1996) Neural image enhancement allows honeybees to see at night. Proc R Soc Lond B Biol Sci 263:1521–1526CrossRefGoogle Scholar
  46. Waser NM (1986) Flower constancy—definition, cause, and measurement. Am Nat 127:593–603CrossRefGoogle Scholar
  47. Wei CA, Dyer FC (2009) Investing in learning: why do honeybees, Apis mellifera, vary the durations of learning flights? Anim Behav 77:1165–1177CrossRefGoogle Scholar
  48. Wei CA, Rafalko SL, Dyer FC (2002) Deciding to learn: modulation of learning flights in honeybees, Apis mellifera. J Comp Physiol Neuroethol Sens Neural Behav Physiol 188:725–737CrossRefGoogle Scholar
  49. Wolf TJ, Schmidhempel P, Ellington CP, Stevenson RD (1989) Physiological correlates of foraging efforts in honeybees—oxygen-consumption and nectar load. Funct Ecol 3:417–424CrossRefGoogle Scholar
  50. Wolf TJ, Ellington CP, Begley IS (1999) Foraging casts in bumblebees: field conditions cause large individual differences. Insect Soc 46:291–295CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2010

Authors and Affiliations

  • John M. Townsend-Mehler
    • 1
    Email author
  • Fred C. Dyer
    • 1
  • Kim Maida
    • 1
  1. 1.Department of ZoologyMichigan State UniversityEast LansingUSA

Personalised recommendations