Localization, morphology, anatomy and ultrastructure of osmophores in species of Rhamnaceae.

Although the presence of scent was described for several species of Rhamnaceae, localization, morphology and structure of osmophores were unknown. We studied different species of the tribes Rhamneae (Rhamnoids clade), Pomaderreae, Colletieae, Paliureae (Ziziphoids clade) and the species Alphitonia excelsa (unknown tribe, Ziziphoids clade). We expect to have a better comprehension of these structures and provide information on which morphological and anatomical characters may support the phylogeny of the family. We localized the osmophores in the margins and top of the sepals using neutral red. Histochemical tests were made on transverse hand-cut sections of fresh sepals. Observations were made with stereoscopic and bright field microscopes, scanning and transmission electron microscopes. Papillae were observed in the zones with positive reaction to reagents. Different kinds of hairs are present in the sepal epidermis besides papillae. Epidermal cells present a striate cuticle with canals and cavities. Druses are abundant in most species. The ultrastructure of epidermal and subepidermal cells shows high metabolic activity: there are vesicles, mitochondria, endoplasmic reticulum, dictyosomes, plastids with lipids and starch. The vascularization is well developed and reaches the top of the sepal where the principal area of volatile components production is localized. The location and abundance of papillae are the most important traits that allow us recognize and characterize the osmophores in Rhamnaceae. There are no clear anatomical and morphological features exclusive of one clade or tribe. Therefore, in contrast to other sporophytic structures of this family, osmophores do not seem to have any systematic value.