Torus Structure and Development in Species of Daphne

Abstract

A torus is present in intervascular pit membranes in the wood of Daphne odora and D. cneorum, but not in D. mezereum. In the two former species, each torus is surrounded by a margo consisting of fibrillar material in a tightly woven pattern. Tori are of greater diameter than pit apertures and completely occlude the apertures during aspiration. Evidence from D. odora indicates that torus deposition is spatially associated with vesicles and a plexus of microtubules, and does not begin until pit border formation is complete. The material deposited during torus synthesis also impregnates the wall of the pre-existing pit membrane. The plasmalemma often is closely appressed to the pit membrane at the site of the developing torus. In half-bordered pit pairs between tracheary elements and parenchyma cells, a torus thickening is deposited only on the side of the tracheary element. As in Osmanthus americanus, it is hypothesised that the presence of tori in species of Daphne prevents rupture of the pit membrane during aspiration.