Abstract
Background and Aims
The functional morphology of Salvia pratensis  flowers was re-investigated, after new insights revealed that pollen  dispensing is one of the main functions of the staminal lever. In  particular, no detailed information was available regarding the process  of pollen transfer and the forces arising between the pollen-bearing  thecae and the pollinating bee's body. The assumption was made that  these forces play a significant role in pollen dispensing.
Methods
The functional morphology of S. pratensis flowers and the interaction between flowers and bees (Apis mellifera)  were studied by reconstructing stress and strains by using qualitative  and semi-quantitative theoretical analysis. Flowers were manipulated to  study the spatial arrangement of the filament and lever, and of the head  and proboscis of the visiting bee inside the tube. Photographs and  films of bee visits on flowers were used to analyse the interaction of  pollinator and staminal lever.
Key Results
The spoon-shaped lower lever of S. pratensis  has a small hole through which a bee introduces its proboscis into the  corolla tube. Although mentioned for the first time by Kerner von  Marilaun in 1891, presented here is the first drawing and the first  photograph showing this interaction in detail. The analysis of the  interaction of flower visitor and the lever mechanism revealed that the  position of bees on different flowers is spatially very similar. Flower  morphology constrains postures of legitimately nectar-probing bees  within narrow bounds. A theoretical discussion on structural elements  and force progression in the flower allows the principles of lightweight  architecture in flower morphology to be recognized.
Conclusions
The staminal lever of S. pratensis  is a pollen-dispensing device. It seems to influence the amount of  pollen deposited on pollinators by determining the forces arising  between the pollinator and the pollen. The relevant forces occur either  during the first, dynamic phase or during the second, almost static  phase of a flower visit.
Key words: Flower–pollinator interaction, bee, Apis mellifera, pollination, pollen uptake, see-saw mechanism, biomechanics, pollen dispensing


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