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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.

Acknowledgments

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.

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