Distribution of vessel size, vessel density and xylem conducting efficiency within a crown of silver birch (Betula pendula)

Abstract

Spatial patterns in vessel diameter, vessel density and xylem conducting efficiency within a crown were examined in closed-canopy trees of silver birch (Betula pendula). The variation in anatomical and hydraulic characteristics of branches was considered from three perspectives: vertically within a crown (lower, middle and upper crown), radially along main branches (proximal, middle and distal part), and with respect to branch orders (first-, second- and third-order branches). Hydraulically weighted mean diameter of vessels (D h) and theoretical specific conductivity of the xylem (k t) exhibited no vertical trend within the tree crown, whereas leaf-specific conductivity of the xylem (LSCt) decreased acropetally. Variation in LSCt was governed by sapwood area to leaf area ratio (Huber value) rather than by changes in xylem anatomy. The acropetal increase in soil-to-leaf conductance (G T) within the birch canopy is attributable to longer path length within the lower-crown branches and higher hydraulic resistance of the shade leaves. D h, k t and LSCt decreased, while vessel density (VD) and relative area of vessel lumina (VA) increased distally along main branches. A strong negative relationship between vessel diameter and VD implies a trade-off between hydraulic efficiency and mechanical stability of xylem. D h and VD combined explained 85.4% of the total variation of k t in the regression model applied to the whole data set. Xylem in fast-growing branches (primary branches) had greater area of vessel lumina per unit cross-sectional area of sapwood, resulting in a positive relationship between branch radial growth rate and k t. D h, k t and LSCt decreased, whereas VD increased with increasing branch order. This pattern promotes the hydraulic dominance of primary branches over the secondary branches and their dominance over tertiary branches. In this way crown architecture contributes to preferential water flow along the main axes, potentially providing better water supply for the branch apical bud and foliage located in the outer, better-insolated part of the crown.