Cell wall thickenings and tylosoid: developmental morphology reveals novelties for secretory canals in Protium ovatum (Burseraceae)

Abstract

We investigated the structure, histochemistry, and ultrastructure of the secretory canals in the vegetative axis of Protium ovatum from a developmental perspective. Samples of roots, stems, and leaves were analyzed using light and transmission electron microscopy. Secretory canals composed of a uniseriate epithelium and a wide lumen occurred in the phloem of all analyzed organs. Schizogenesis and lysigenesis were merging processes involved in the origin, growth, ramification, and fusion of the secretory canals, forming an anastomosed secretory net. Essential oils, polysaccharides, and proteins were detected in the epithelial cells, as well as plastids with poorly developed thylakoids, dictyosomes, rough endoplasmic reticulum, polysomes, and oil drops, showing the mixed nature of the secretion. Epithelial cells exhibited pectin-cellulosic thickenings in the anticlinal and radial walls. These thickenings may act in directing the secretion flux toward the lumen, protecting the neighboring tissues from the toxicity of secreted metabolites. Structural irregularities observed in the mitochondria cristae in epithelial cells may be associated with processes induced by toxic substances. Epithelial cells protruded into the lumen and became lignified in the outer portion of the secondary phloem, obliterating the non-functional secretory canals. We propose that this phenomenon presents a physiological significance similar to that of tylose, preserving the secretion flow inside the active portions of the secretory system. To our knowledge, epithelial cells with wall thickenings, mitochondria with structural abnormalities, and obliteration of non-functional canals are features reported for the first time for Burseraceae. These features have important functional significance for Burseraceae secretory system and contribute to a deeper knowledge of P. ovatum, a medically and economically important plant.