ECCENTRIC GROWTH AND GROWTH STRESS IN INCLINED STEMS OF GNETUM GNEMON

Gnetum gnemon L. (Gnetales) forms hardwood-like secondary xylem in its trunks and branches although it is a gymnosperm. The present study tested the surface growth stress in relation to anatomical and chemical properties of the secondary xylem in inclined and vertical stems of G. gnemon using morphological and chemical composition analyses. Secondary growth was promoted on the upper half of the cross section in an inclined stem; at the same time, tensile growth stress increased on the upper side and decreased on the lower side of the inclined stem. However, formation of reaction wood fibers was not detected on either side. The microfibril angle was associated with differences in tensile growth stress. Thus, we conclude that negative gravitropism in G. gnemon is caused by a synergistic effect of increased tensile growth stress as well as the promotion of secondary growth on the upper side of the inclined stem. Our results are comparable to the negative gravitropism observed in the family Magnoliaceae, which does not form gelatinous fibers in its tension wood.