Colleters in Bathysa cuspidata (Rubiaceae): Development, ultrastructure and chemical composition of the secretion

This paper describes the development of colleters of Bathysa cuspidata, Rubiaceae, considering anatomical, histochemical and ultrastructural aspects and going from first differentiation stages until senescence. Further, the chemical composition of the secretion is investigated. The samples were prepared according to the usual techniques for light microscopy and scanning and transmission electron microscopy. Electrophoresis and thin layer chromatography (TLC) were used to confirm the results obtained in the histochemical tests. The colleters occur at the ventral surface of the stipules which protect the leaf primordia as well as the shoot meristem. The origin of the colleters is mixed, involving protoderm and ground meristem. The Bathysa colleters are of the standard type or are bifurcated; this latter type is documented here for the first time for Rubiaceae. Colleter secretion is a mucilage rich in protein, as determined by histochemical tests and confirmed by chemical analysis. Phenolic compounds and terpenes were detected only in the colleters themselves, but not in the secretion. The epithelial cells present conspicuous nuclei and nucleoli and the cytoplasm is rich in dictyosomes, endoplasmic reticulum, mitochondria, vesicles and small vacuoles with a fibrillar content. The accumulation of phenolic compounds and terpenes, the formation of a large central vacuole, the increase of the intercellular and subcuticular spaces occupied by the secretion and, finally, the darkening and the wilting of the colleters characterize the senescence of these structures. The secretion process of the colleters of B. cuspidata suggests a process of programmed cell death.