Structure and ontogeny of the fissured stems of Callaeum (Malpighiaceae)

Abstract

Stem ontogeny and structure of two neotropical twining vines of the genus Callaeum are described. Secondary growth in Callaeum begins with a typical regular cambium that gradually becomes lobed as a result of variation in xylem and phloem production rates in certain portions of the stem aligned with stem orthostichies. As development progresses, lignified ray cells of the initially formed secondary xylem detach on one side from the adjacent tissues, forming a natural fracture that induces the proliferation of both ray and axial nonlignified parenchyma. At the same time, parenchyma proliferation takes place around the pith margin and generates a ring of radially arranged parenchyma cells. The parenchyma generated in this process (here termed disruptive parenchyma) keeps dividing throughout stem development. As growth continues, the parenchyma finally cleaves the lignified axial parts of the vascular system into several isolated fragments of different sizes. Each fragment consists of xylem, phloem and vascular cambium and is immersed in a ground matrix of disruptive parenchyma. The cambium present in each fragment divides anticlinally to almost encircle each entire fragment and maintains its regular activity by producing xylem to the centre of the fragment and phloem to the periphery. Additionally, new cambia arise within the disruptive parenchyma and produce xylem and phloem in various polarities, such as xylem to the inside and phloem to the outside of the stem, or perpendicularly to the original cambium. Unlike the very distinctive stem anatomical architecture resulting from this cambial variant in Callaeum, its secondary xylem and phloem exhibit features typical of lianas. These features include very wide conducting cells, abundant axial parenchyma, high and heterocellular rays and gelatinous fibres.