Carpellary appendages in Nymphaea and Victoria (Nymphaeaceae): evidence of their role as osmophores based on morphology, anatomy and ultrastructure

In flowers of Nymphaea and Victoria, carpellary appendages are regarded as structures related to pollination by deceit of night-blooming species. In this study, the anatomy, histochemistry and ultrastructure of carpellary appendages were analysed to investigate their possible role in the production of volatile compounds in nocturnal species Nymphaea amazonum, N. gardneriana, N. prolifera (Nymphaea subgenus Hydrocallis) and Victoria cruziana, and in diurnal species N. caerulea (Nymphaea subgenus Brachyceras). Carpellary appendages were studied using light microscopy and scanning and transmission electron microscopy from pre-anthesis to the second day of anthesis. Anatomical and ultrastructural features are characteristic of osmophores. In all species, the most frequent components in secretory cells are amyloplasts, lipid bodies, mitochondria, rough endoplasmic reticulum and elaioplasts. The epidermis and multilayered parenchyma accumulate abundant starch grains and lipophilic substances, both of which vanish during anthesis. Amorphous substances are deposited between the plasmalemma and the outer cell wall of epidermal cells, and are then released by cuticular diffusion. Odour production in carpellary appendages might be an ancient role of primary importance both in diurnal and nocturnal species that are pollinated by deceit. Olfactory and visual cues of small carpellary appendages in Nymphaea subgenus Brachyceras correspond to bee pollination, and large carpellary appendages in subgenus Hydrocallis and Victoria represent parallel functional specializations of the flowers to the attraction and reward for exclusive beetle pollination.