Osmophores of Caryocar brasiliense (Caryocaraceae): a particular structure of the androecium that releases an unusual scent.

Caryocar brasiliense is a flagship species of the Brazilian Cerrado. It produces flowers with a strong peculiar scent, which are pollinated by bats and occasionally moths with short mouthparts. However, the cues responsible for attracting these nocturnal pollinators remain unknown. We aimed to identify osmophores of C. brasiliense, describe the ultrastructure of the cells involved in the synthesis and release of floral odour, and identify the constituents of the floral bouquet. We performed field observations and histochemical and ultrastructural analyses of flowers focusing on the androecium. Gas chromatography-mass spectrometry was used to analyse the scents emitted. Filament epidermal cells were found to possess an unusual shape and be responsible for the main production and release of odour. These cells, called foraminous cells, are elongate and possess pores where their cell walls are abruptly thin. The cuticle is practically absent over the pores, which facilitates odour emission. The foraminous cells have conspicuous nuclei and organelle-rich cytoplasm where oil droplets can be seen prior to anthesis. The features of these cells remain similar during anthesis, but many vesicles fuse with the plasma membrane and the number of oil droplets in the cytosol decreases. Twenty-nine components were found in the scent, especially fatty acid derivatives and N- and S-bearing compounds. Our analyses revealed that the androecium of C. brasiliense has a particular structure that acts as an osmophore. The scent from the androecium resembles that of the entire flower, which is an unprecedented finding for a plant with single flowers as the pollination unit.