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Immunocytochemical Evidence that Secretion of Pectin Occurs During Gel (Gum) and Tylosis Formation in Trees

First, third, and fourth authors: Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, P.O. Box 3800, Sainte-Foy, Quebec G1V 4C7, Canada; and second author: ORSTOM, Institut Français de Recherche Scientifique pour le Développement en Coopération, Laboratoire de Phytopathologie, B.P. 5045, 34032 Montpellier, France

During gel (gum) formation in angiosperm trees, fibrillar material accumulated in protective layers of xylem parenchyma cells before being secreted across half-bordered pit membranes into vessel elements. Immunogold labeling demonstrated that this fibrillar material was mainly composed of partially esterified pectic polysaccharides. The primary wall of expanding tyloses, an extension of the parenchyma protective layer, secreted similar pectic substances to completely block vessel elements. In most studies, these occluding structures were reported to be formed in response to causative factors such as aging processes, injuries, or infections. Current observations support the view that partial to complete embolism, which almost always accompanies these factors, might be the main cause triggering the formation of vessel occlusions. Whereas pectin seems to be the basic component of gels (gums) and of the external layer of tyloses, other substances, such as phenols, were also detected either as a part of these plugs or as accumulations beside them in vessels. Finally, it is proposed that the term ‘gel’ instead of ‘gum’ be used in future studies to describe the occluding material secreted by ray and paratracheal parenchyma cells.