Buzz Pollination

Pollinators frequently use complex motor routines to find and extract floral rewards. Studies of polli-nators foraging for nectar rewards indicate these routines are typically learned, and that constraints associated with learning and memory give pollinators incentive to continue foraging on these flowers. However, plants offer rewards besides nectar, including pollen, lipids and essential oils. In particular, bees use a complex motor routine termed floral sonication to extract pollen, their primary source of protein, from the more than 6% of flowering plant species (>22 000 species) that conceal pollen rewards within tube-like poricidal anthers. If floral sonication requires learning, this pollen extraction behaviour could contribute to floral fidelity. However, no studies have quantified the effect of experience on flower handling for bees extracting pollen from poricidal species. We therefore examined the degree to which floral sonication behaviour was modified by experience. We found that the key elements of the soni-cation motor routine appeared in full-blown form in a flower-naïve bee's first visit to a flower. We additionally found consistent, albeit modest, effects of experience on certain aspects of sonication behaviour. The latency to sonicate slightly decreased with experience. Bees also adjusted the length and amplitude of their sonication buzzes in response to pollen receipt. We conclude that the role of experience in foraging for concealed pollen rewards is different from that reported for nectar rewards. We offer an alternative explanation for its function in sonication. Finally, we discuss alternative hypotheses for the function of poricidal anthers and for how pollen-bearing plants may ensure floral fidelity even in the absence of a significant impact of experience on pollen extraction behaviour.

Heteranthery is thought to reflect a division of labor, with some anthers serving a pollinator-feeding function and others serving a pollinating function. Mutualism theory predicts that each participant should try to maximize the benefit it receives from its partner: plants should allocate more pollen to pollination, and pollinators should collect more pollen. Accordingly, plant and pollinator may engage in a ‘tug of war’ with respect to pollen from each anther type, resulting in incomplete division of labor. Here, we explored this idea by conducting a fully factorial manipulation of the availability of pollen in long and short anthers of staminate flowers of Solanum houstonii. We found the following: (1) Bumble bees (Bombus impatiens) preferred to sonicate (collect pollen from) short anthers over long anthers, consistent with a role as feeding and pollinating anthers, respectively; (2) Blocking short anther pores alone increased sonication of long anthers and resulted in collection of pollen from long anthers; (3) Blocking long anther pores alone did not influence sonication of short anthers; (4) The increase in sonication of long anthers, when short anther pores are blocked, was greater when pollen was available in long anthers; (5) Despite shifting sonication effort to long anthers, bees do not move their bodies closer to long anther pores where pollen could be collected more effectively; and (6) analysis of the growth of corbicular loads over time spent buzzing indicates that significant amounts of pollen are collected from long anthers as well as short anthers. We conclude that bees can flexibly increase pollen collection from pollinating anthers, but are constrained from fully exploiting this pollen. This results in checks and balances between plant and bee that may help maintain heteranthery.