Transpiration-induced axial and radial tension gradients in trunks of Douglas-fir trees.

We determined the axial and radial xylem tension gradients in trunks of young Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Axial specific conductivity (k(s-a)) and sap flux density (Js) were measured at four consecutive depths within the sapwood at a stem height of 1 m. By definition, at a given position in the bole, Js is a function not only of k(s-a) but also of the driving force for water movement. The Js:k(s-a) ratio was therefore used to estimate axial tension gradients and the radial gradients at a stem height of 1 m were calculated from the differences in axial tension gradients at each depth. Tracheid lumen diameter and tracheid length were used to predict differences in k(s-a) and its divergence from the theoretical k(s-a) determined by the Hagen Poisseuille equation. The ratio of k(s-a) (determined in the laboratory) to Js (measured in the field) varied with depth in the sapwood, resulting in non-uniform axial and radial tension gradients from inner to outer sapwood. Transpiration-induced axial tension gradients were in the range of 0.006-0.01 MPa m(-1) excluding the gravitational tension gradient. At a stem height of 1 m, radial tension gradients were in the range of 0.15-0.25 MPa m(-1) and were lower in the middle sapwood than in the inner or outer sapwood. Axial tension gradients were 44-50% higher in the outer sapwood than in the inner sapwood. At a stem height of 1 m, radial Js, calculated on the basis of radial tension gradients and measured radial specific conductivity (k(s-r)), was about two orders of magnitude smaller than axial Js. Our findings indicate that large radial tension gradients occur in the sapwood and clarify the role played by xylem k(s-a) and k(s-r) in determining in situ partitioning of Js in the axial and radial directions.

Age- and position-related changes in hydraulic versus mechanical dysfunction of xylem: inferring the design criteria for Douglas-fir wood structure.

We do not know why trees exhibit changes in wood characteristics as a function of cambial age. In part, the answer may lie in the existence of a tradeoff between hydraulic properties and mechanical support. In conifers, longitudinal tracheids represent 92% of the cells comprising the wood and are involved in both water transport and mechanical support. We used three hydraulic parameters to estimate hydraulic safety factors at several vertical and radial locations in the trunk and branches: vulnerability to cavitation; variation in xylem water potential (psi); and xylem relative water content. The hydraulic safety factors for 12 and 88 percent loss of conductivity (S(H12) and S(H88), representing the hydraulic safety factors for the air entry point and full embolism point, respectively) were determined. We also estimated the mechanical safety factor for maximum tree height and for buckling. We estimated the dimensionless hydraulic and mechanical safety factors for six seedlings (4 years old), six saplings (10 years old) and six mature trees (> 110 years old) of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco). Over the natural range of psi, S(H12) decreased linearly from treetop to a minimum of 0.95 at the tree base. Young and mature trees had S(H12) values 1.4 and 1.3 times higher, respectively, at their tips (juvenile wood) than at their bases (mature wood). Modeling analyses indicated that if trees were made entirely of mature wood, S(H12) at the stem base would be only 0.7. The mechanical safety factor was 1.2 times higher for the base of the tree than for the rest of the tree. The minimum mechanical safety factor-1.6 for the critical buckling height and 2.2 for the critical buckling load-occurred at the base of the live crown. Modeling analysis indicated that if trees were made only of mature wood, these values would increase to 1.7 and 2.3, respectively. Hydraulic safety factors had values that were less than half those for mechanical safety factors, suggesting that wood structure in Douglas-fir has evolved primarily as a result of selection for hydraulic safety rather than mechanical safety. The results suggest that forest managers must consider the role of juvenile wood in tree physiology to avoid producing plantations vulnerable to drought.

Dynamics of water transport and storage in conifers studied with deuterium and heat tracing techniques.

The volume and complexity of their vascular systems make the dynamics of long-distance water transport in large trees difficult to study. We used heat and deuterated water (D2)) as tracers to characterize whole-tree water transport and storage properties in individual trees belonging to the coniferous species Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. The trees used in this study spanned a broad range of height (13.5-58 m) and diameter (0.14-1.43 m). Sap flow was monitored continuously with heat dissipation probes near the base of the trunk prior to, during and following injection of D2O. The transit time for D2O transport from the base of the trunk to the upper crown and the tracer residence time were determined by measuring hydrogen isotope ratios in water extracted from leaves sampled at regular intervals. Transit times for arrival of D2O in the upper crown ranged from 2.5 to 21 d and residence times ranged from 36 to 79 d. Estimates of maximum sap velocity derived from tracer transit times and path length ranged from 2.4 to 5.4 m d(-1). Tracer residence time and half-life increased as tree diameter increased, independent of species. Species-independent scaling of tracer velocity with sapwood-specific conductivity was also observed. When data from this study were combined with similar data from an earlier study of four tropical angiosperm trees, species-independent scaling of tracer velocity and residence time with sapwood hydraulic capacitance was observed. Sapwood capacitance is an intrinsic tissue-level property that appears to govern whole-tree water transport in a similar manner among both tracheid- and vessel-bearing species.

Stem respiratory potential in six softwood and four hardwood tree species in the central cascades of Oregon.

Mature and old growth trees of varying sapwood thickness were compared with regard to stem respiration. An increment core-based, laboratory method under controlled temperature was used to measure tissue-level respiration (termed respiratory potential) of ten different tree species. Bark (dead outer and live inner combined), sapwood, and heartwood thickness measurements were used to predict sapwood volume from stem diameter (including bark) for four of the ten species. These predictions of sapwood volume were used to scale respiratory potential to the main-bole level (excluding all branches). On the core level, species that maintained narrow sapwood (8-16% of bole radius) such as Pseudotusga menziesii, Taxus brevifolia, and Thuja plicata, had sapwood respiratory potentials in the lower bole that were 50% higher (P<0.05) than species with wide sapwood (>16% of bole radius), such as Abies amabilis, Pinus monticola, and Tsuga heterophylla. This pattern was not observed for inner bark respiratory potential, or for sapwood respiratory potential within the crown. On the main-bole level, respiratory potential per unit volume was inversely correlated to the live bole volumetric fraction (inner bark plus sapwood divided by whole bole volume) (Adj. R(2)=0.6). Specifically, tree species with 18-20% of the main bole alive potentially respired 1.3-3 times more per unit live bole volume than species with over 40%, suggesting that the live bole was less metabolically active in tree species that maintained large volumes of sapwood.

Shoot and root vulnerability to xylem cavitation in four populations of Douglas-fir seedlings.

The objectives of this study were to assess the range of genotypic variation in the vulnerability of the shoot and root xylem of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings to water-stress-induced cavitation, and to assess the trade-off between vulnerability to cavitation and conductivity per unit of stem cross-sectional area (k(s)), both within a species and within an individual tree. Douglas-fir occupies a broad range of environments and exhibits considerable genetic variation for growth, morphology, and drought hardiness. We chose two populations from each of two varieties (the coastal var. menziesii and the interior var. glauca) to represent environmental extremes of the species. Vulnerability curves were constructed for shoots and roots by plotting the percentage loss in conductivity versus water potential. Vulnerability in shoot and root xylem varied genetically with source climate. Stem xylem differed in vulnerability to cavitation between populations; the most mesic population, coastal wet (CW), was the most susceptible of the four populations. In the roots, the most vulnerable population was again CW; the interior wet (IW) population was moderately susceptible compared with the two dry populations, coastal dry (CD) and interior dry (ID). Root xylem was more susceptible to cavitation than stem xylem and had significantly greater k(s). The trade-off between vulnerability to cavitation and k(s), however, was not evident across populations. The most vulnerable population (CW) had a shoot k(s) of 0.534 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1), compared with 0.734 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1) for the less vulnerable CD stems. In the roots, IW was more vulnerable than ID, but had the same k(s).