EARLYWOOD VESSELS IN RING-POROUS TREES BECOME FUNCTIONAL FOR WATER TRANSPORT AFTER BUD BURST AND BEFORE THE MATURATION OF THE CURRENT-YEAR LEAVES

Abstract

This paper reviews the development of xylem vessels in ring-porous dicots and the corresponding leaf phenology. Also included are our original observations on the time-course of vessel element growth, secondary wall deposition, and end wall perforation in the deciduous hardwood Kalopanax septemlobus. Different patterns of xylem growth and phenology serve different strategies of the species for adaptation to seasonal climates. Trees with ring-porous xylem form wide earlywood vessels (EWV) in spring and narrow latewood vessels in summer. The wide EWV become embolized or blocked with tyloses by the end of the growing season while the narrow vessels may remain functional for many years. The co-occurrence of wide and narrow vessels provides both efficiency and safety of the water transport as well as a potentially longer growing season. It has for a long time been assumed that EWV in ring-porous hardwoods are formed in early spring before bud burst in order to supply sap to growing leaves and shoots. However, the full time-course of development of EWV elements from initiation of growth until maturation for water transport has not been adequately studied until recently. Our observations clarify a crucial relationship between leaf maturation and the maturation of earlywood vessels for sap transport. Accumulated new evidence shows that EWV in branches and upper stem parts develop earlier than EWV lower in the stem. The first EWV elements are fully expanded with differentiated secondary walls by the time of bud burst. In lower stem parts, perforations in vessel end walls are formed after bud burst and before the new leaves have achieved full size. Therefore, the current-year EWV network becomes functional for water transport only by the time when the first new leaves are mature.