The effects of cambial age and position within the stem on specific conductivity in Douglas-fir ( Pseudotsuga menziesii) sapwood

Abstract

Specific conductivity (k s, m2s–1MPa–1) describes the permeability of xylem and is determined by all aspects of xylem anatomy that create resistance to the flow of water. Here we test the hypothesis that k s is a function of radial and vertical position within the stem, rather than solely a function of cambial age (ring number from the pith), by measuring k s on samples excised from 35-year-old Douglas-fir [Pseudotsuga menziesii var. menziesii (Mirb.) Franco] trees at six heights and two or three radial positions. Sapwood k s decreased from the cambium to the heartwood boundary, and the difference between outer and inner sapwood increased with height in the tree. Beneath the live crown, inner sapwood had 80–90% the k s of outer sapwood, but only 55% just 10 m higher in the stem (about 10 nodes down from the tree top). Outer sapwood k s peaked near the base of the crown and declined toward both the base and top of the stem. These patterns can be explained by two superimposed effects: the effect of cambial age on the dimensions of tracheids as they are produced, and the effect of xylem aging, which may include accumulation of emboli and aspiration of bordered pits. Tracheid lumen diameter and earlywood and latewood density and width, all factors known to vary with cambial age, were measured on different trees of the same age and from the same stand. Lumen diameter increased with cambial age, whereas the proportion of latewood and growth ring density increased after an initial decrease in the first 5 years. Our results suggest that the effect of cambial age on xylem anatomy is not sufficient to explain variation in k s. Instead, physical position (both vertical and radial) in the stem and cambial age must be considered as determinants of conductivity.