TYLOSES AND GUMS: A REVIEW OF STRUCTURE, FUNCTION AND OCCURRENCE OF VESSEL OCCLUSIONS

Vessel occlusion through tyloses or gums is a natural phenomenon occurring with aging and heartwood formation, and in sapwood in response to vessel embolism. These types of vessel occlusion play a crucial role to limit the spread of pathogens and wood decay organisms, also as part of compartmentalization after wounding. In the sapwood, they can be considered to be an effective stress response. Here we review the literature on tyloses and gums in hardwoods, starting with the detailed 19th century account on tyloses by Hermine von Reichenbach. The structural diversity of tyloses (from thin-walled to sclerotic) and gum deposits is highlighted and illustrated. Our understanding of the development of vessel occlusions through vessel contact cells of the ray and axial parenchyma has greatly increased over the last decades, also thanks to ultrastructural and immunocytological studies. We critically discuss the postulated relationships between vessel-to-ray parenchyma pit size and vessel size and the incidence of either tyloses or gums and review the occurrence of these types of vessel occlusions in extant and fossil dicots. All factors identified in the literature as stimuli for vessel occlusion probably act through vessel embolism as a single direct trigger. Attempts in the literature to relate vessel occlusion with mechanisms of vessel refilling and embolism repair are controversial and invite more experimental research.