to gains is known for human beings (Tversky & Kahneman 1981). This suggests that in a competitive market for pollinator service, there is a decelerating payoff to plants (in terms of attracting pollinators) for increasing the quality of their nectar. The same likely applies to volume. We also found that losses (declining concentration) and energetic costs are perceptually weighted relative to gains. Concerning the latter, a change in floral morphology that increases or decreases handling cost only slightly can have important effects on the attractiveness of the flower to pollinators. I think that we can expect these patterns of behavior to apply generally to other pollen- and nectar-foraging taxa.
The perceptual scale based on the honeybee dance successfully predicted, in laboratory studies using artificial flowers, choice behavior in relation to expected sucrose concentration and variance in sucrose concentration. Preference for concentration depends on the magnitude of difference between the concentrations and magnitudes of the concentrations. Response to variance in our experiments depended on the magnitude of the expected concentrations and of the magnitude in variance of the high-variance flower. Choices among flowers differing in expected nectar volume and variance in volume, and pollen quality, all are explained by non-linear perceptual relationships; however, there is variation among bees in their perception. Our studies show that genotypic variation is one source of variation in perception. Genotypic variation in honeybees' choice behavior is consistent with their differing perceptual scales indicated by the dance. Finally, the proboscis extension response; (PER) is just beginning to be used to study perception of sucrose concentration. Thus far, PER results suggest genotypic variation in response and experiential factors which modulate the response; PER also has promise for studying risk-sensitive foraging behavior (Shafir et al. 1999). This technique will be especially useful for exploring perceptual differences among individual bees.
I thank Dr Peter Vanderborght of the International Society for Horticultural Science for giving me permission to reprint parts of Waddington (1997) here. I also thank Academic Press and Blackwell Wissenschafts-Verlag for permitting me to reproduce figures. Dr C. Mindy Nelson made helpful comments on the manuscript.
Banschbach VS & Waddington KD (1994) Risk-sensitive foraging in honeybees: no consensus among individuals and no effect of colony honey stores. Anim Behav 7:933-941
Barth FG (1985) Insects and flowers: the biology of a partnership. Princeton University Press, Princeton, NJ
Bitterman ME, Menzel R, Fietz A & Schafer S (1983) Classical conditioning of proboscis extension in honeybees (Apis mellifera). J Comp Psychol 97:107-119 Buchmann SL & Cane JH (1989) Bees assess pollen returns while sonicating Solanum flowers. Oecologia 81:289-294 Calderone N & Page RE (1988) Genotypic variability in age polyethism and task specialization in the honeybee, Apis mellifera (Hymenoptera; Apidae). BehavEcol Sociobiol 22:17-25
Calderone N & Page RE (1992) Effects of interactions among genetically diverse nestmates on task specialization by foraging honeybees (Apis mellifera). Behav Ecol Sociobiol 30:219-226
Caraco T (1980) On foraging time allocation in a stochastic environment. Ecology 61:119-128
Carterette EC & Friedman MP (1974) Handbook ofperception, vol. 2. Academic Press, New York
Charnov E (1976) Optimal foraging: attack strategy of a mantid. Am Nat 110:141-151 Deng G (1996) Foraging performance of honeybees (Apis mellifera). PhD thesis,
University of Miami, FL Faegri K & van der Pijl, L (1979) The principles of pollination ecology, 3rd edn. Pergamon Press, New York
Fewell JH & Page RE (1993) Genotypic variation in foraging responses to environmental stimuli by honeybees, Apis mellifera. Experientia 49:1106-1112 Frisch K von (1967) Dance language and orientation of bees. Belknap Press, Cambridge, MA Grant V (1949) Pollination systems as isolating mechanisms in flowering plants. Evolution 3:82-97
Greggers U & Menzel R (1993) Memory dynamics and foraging strategies of honeybees.
Behav Ecol Sociobiol 32:17-29 Guzman-Novoa E & Gary NE (1993) Genotypic variability of components of foraging behaviour in honeybees (Hymenoptera; Apidae) J Econ Entomol 86:715-721 Harder LD (1990) Behavioral responses by bumble bees to variation in pollen availability. Oecologia 85:41-47 Harder L & Real LA (1987) Why are bumble bees risk averse? Ecology 68:1104-1108 Kacelnik A & Bateson M (1996) Risky theories - the effects of variance on foraging decisions. Am Zool 36:402-434 Macior LW (1970) The pollination ecology of Pedicularis in Colorado. Am J Bot 57:716-728 Milne CP, Hellmich RL & Pries KJ (1986) Corbicular size in workers from honeybee lines selected for high or low pollen hoarding. J Apic Res 25:50-52 Page RE, Waddington KD, Hunt GJ and Fondrk MK (1995) Genetic determinants of honeybee foraging behaviour. Anim Behav 50:1617-1625 Page RE, Erber J & Fondrk MK (1998) The effect of genotype on response thresholds to sucrose and foraging behavior of honeybees (Apis mellifera L.). J Comp Physiol A 182:489-500
Perez SP & Waddington KD (1996) Carpenter bee (Xylocopa micans) risk-indifference and a review of nectarivore risk-sensitivity studies. Am Zool 36:435-446 Possingham HP, Houston AI & McNamara JM (1990) Risk-averse foraging in bees: a comment on the model of Harder and Real. Ecology 71:1622-1624 Pyke GH (1979) Optimal foraging in bumblebees: rule of movement between flowers within inflorescences. Anim Behav 27:1167-1181 Pyke GH (1984) Optimal foraging theory: a critical review. Ann RevEcol Syst 15:523-575 Rachlin H (1989) Judgement, decision, and choice. WH Freeman, New York Rasheed SA & Harder LD (1997a) Foraging currencies for non-energetic resources:
pollen collection by bumblebees. Anim Behav 54:911-926 Rasheed SA & Harder LD (1997b) Economic motivation for plant species preferences of pollen-collecting bumble bees. Ecol Entomol 22: 209-219 Raveret-Richter M & Waddington KD (1993) Past foraging experience influences honeybee dance behaviour. Anim Behav 46:123-128 Real L, Ott J & Silverfine E (1982) On the trade-off between the mean and the variance in foraging: effects of spatial distribution and color preference. Ecology 63:1617-1623
Real LA, Ellner S & Harder LD (1990) Short-term energy maximization and risk-
aversion in bumblebees: comments on Possingham et al. Ecology 71:1625-1628 Schmid-Hempel P, Kacelnik A & Houston AI (1985) Honey bees maximize efficiency by not filling their crop. Behav Ecol Sociobiol 17:61-66 Schmidt JO (1982) Pollen foraging preferences of honeybees. Southwest Nat 7:255-259 Schmidt JO (1984) Feeding preferences of Apis mellifera L. (Hymenoptera: Apidae):
individual versus mixed pollen species. J Kans Entmol Soc 57:323-327 Schmidt JO & Johnson BE (1984) Pollen feeding preference of Apis mellifera, a polylectic bee. Southwest Nat 9:41-47 Seeley T (1986) Social foraging by honeybees: how colonies allocate foragers among patches of flowers. Behav Ecol Sociobiol 19:343-354 Selten R & Shmida A (1991) Pollinator foraging and flower competition in a game equilibrium model. In: Selten R (ed) Game theory in behavioural sciences, pp 195-256. Springer-Verlag, Berlin Shafir S (1994) Intransitivity of preferences in honeybees: support for "comparative"
evaluation of foraging options. Anim Behav 48:55-67 Shafir S, Wiegmann DD, Smith BH & Real L (1999) Risk-sensitive foraging: choice behaviour of honeybees in response to variability in volume of reward. Anim Behav 57:1055-1061
Stephens DW & Charnov E (1982) Optimal foraging: some simple stochastic models.
Behav Ecol Sociobiol 10:251-263 Stephens DW & Krebs JR (1986) Foraging theory. Princeton University Press, Princeton, NJ
Stephens DW & Paton SR (1986) How constant is the constant of risk-aversion? Anim Behav 34:1659-1667
Tversky A & Kahneman D (1981) The framing of decisions and psychology of choice. Science 211:453-458
Waddington KD (1981) Factors influencing pollen flow in bumblebee-pollinated
Delphiniumvirescens. Oikos 37:153-159 Waddington KD (1982) Honeybee foraging profitability and round dance correlates. J Comp Physiol 148:279-301
Waddington KD (1983) Foraging behavior of pollinators. In: Real L (ed) Pollination biology, pp 213-239. Academic Press, NY
Waddington KD (1985) Cost-intake information used in foraging. J Insect Physiol 31:891-897
Waddington KD (1995) Bumblebees do not respond to variance in nectar concentration. Ethology 101:33-38
Waddington KD (1997) Foraging behavior of nectarivores and pollen collectors. Acta Hort 437:175-191
Waddington KD & Gottlieb N (1990) Actual vs. perceived profitability: a study of floral-choice of honeybees. J Insect Behav 3:429-441
Waddington KD & Holden LR (1979) Optimal foraging: on flower selection by bees. Am Nat 114:179-196
Waddington KD & Kirchner W (1992) Acoustical and behavioral correlates of profitability of food sources in honeybee round dances. Ethology 92:1-6
Waddington KD, Allen T & Heinrich B (1981) Floral preferences of bumblebees (Bombus edwardsii) in relation to intermittent versus continuous rewards. Anim Behav
Waddington, KD, Nelson CM & Page RE (1998) Effects of pollen quality and genotype on the dance of foraging honeybees. Anim Behav 56:35-39
Wolf T & Schmid-Hempel P (1990) On the integration of individual foraging strategies with colony ergonomics in social insects: nectar collection in honeybees. Behav Ecol Sociobiol 27:103-111
Was this article helpful?