Stem growth and respiration in loblolly pine plantations differing in soil resource availability

Stem respiration and growth in 10-year-old loblolly pine (Pinus taeda L.) plantations were measured monthly during the third year of fertilization and irrigation treatments to determine whether soil resource availability differentially altered growth and respiration in stem tissue. Fertilized trees had significantly greater stem biomass, stem nitrogen concentration ([N]) and growth rate than unfertilized trees. Stem respiration (Rt) was significantly greater in fertilized trees when expressed on a per unit surface area (Rt,a, micromol CO2 m-2 s-1), sapwood volume (Rt,v, micromol CO2 m-3 s-1), or mass (Rt,w, nmol CO2 g-1 s-1) basis; however, there was no difference between treatments when expressed as a function of stem N content (Rt,n, micromol CO2 (mol N)-1 s-1). Irrigation had no significant effect on Rt or annual stem growth. Daily total respiration (Rd, mol CO2 m-2 day-1) and stem diameter growth both had a seasonal bimodal pattern with peaks in early spring and midsummer. Stem [N] declined significantly during the growing season. Stem growth rate and [N] explained 75% of the seasonal variation in temperature-normalized Rt,a. The mature tissue method was used to partition total stem respiration (Rt) into maintenance (Rm) and growth (Rg) components. There was a linear correlation between winter Rt,v, a measure of basal Rm, and sapwood N content; however, Rt,v per unit N was greater in January before diameter growth started than in the following December after growth ceased, indicating that Rt,v declined as stem diameter increased. Consequently, estimates of annual maintenance respiration (RM) based on January data were 44% higher than estimates based on December data. Growth respiration was correlated with stem growth rate (r2 = 0.55). The growth respiration coefficient (rg)-the slope of the relationship between Rg and stem growth rate-was 0.24. Respiration accounted for 37% of annual stem carbon budget. Stem carbon-use efficiency (CUE)-the ratio of stem growth to stem growth plus respiration-averaged 0.63 and was unaffected by fertilization.     

Gas in stems: abundance and potential consequences for tree biomechanics

Secondary xylem of woody plants has a large volumetric proportion of gas occupying spaces that would otherwise be filled with water. We examined whether these gas-filled voids have a mechanical role by either decreasing the fresh mass the tree must support (by replacing some of the water with gas) or by providing inexpensive filler to increase stem diameter (thereby increasing the second moment of area at the expense of the modulus of elasticity and modulus of rupture). Calculations from published data show that temperate softwood species (n = 26) average 18 and 50% gas by volume for sapwood and heartwood, respectively; temperate hardwood species (n = 31) average 26% gas by volume in both the sapwood and heartwood; and tropical species (n = 52) with mixed sapwood and heartwood have 18% gas by volume. In this paper, we develop equations to show how gas affects the mechanical behavior of tree stems, and describe model results to show how gas affects mechanical stability, based on mass and stem diameters for six 34-year-old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. For the same applied load, modeled stems in which the gas space was filled with water differed in their surface stresses by < 2% from modeled stems in the native state (partially gas-filled), indicating no practical benefit from a reduction in stem mass due to gas. A second modeling scenario compared the native state to stems in which gas was removed and stem diameters decreased (and material properties adjusted to concur with the increased wood density) to conserve mass. Removal of the gas-filled voids resulted in up to 41% higher surface stress for the same applied load, caused by a decrease in the second moment of area greater than the increase in modulus of elasticity. Trees with gas removed had higher modulus of rupture, but could withstand up to 14% lower maximum wind forces than trees in their native state, suggesting a biomechanical role for the gas if the model assumptions are valid. The gas content may, however, have evolved in response to pressures unrelated to biomechanics. We discuss some of its potential effects on sapwood physiology.     

Relationships between stem diameter, sapwood area, leaf area and transpiration in a young mountain ash forest

We examined relationships between stem diameter, sapwood area, leaf area and transpiration in a 15-year-old mountain ash (Eucalyptus regnans F. Muell.) forest containing silver wattle (Acacia dealbata Link.) as a suppressed overstory species and mountain hickory (Acacia frigescens J.H. Willis) as an understory species. Stem diameter explained 93% of the variation in leaf area, 96% of the variation in sapwood area and 88% of the variation in mean daily spring transpiration in 19 mountain ash trees. In seven silver wattle trees, stem diameter explained 87% of the variation in sapwood area but was a poor predictor of the other variables. When transpiration measurements from individual trees were scaled up to a plot basis, using stem diameter values for 164 mountain ash trees and 124 silver wattle trees, mean daily spring transpiration rates of the two species were 2.3 and 0.6 mm day(-1), respectively. The leaf area index of the plot was estimated directly by destructive sampling, and indirectly with an LAI-2000 plant canopy analyzer and by hemispherical canopy photography. All three methods gave similar results.     

Sapwood temperature gradients between lower stems and the crown do not influence estimates of stand-level stem CO(2) efflux

Temperature plays a critical role in the regulation of respiration rates and is often used to scale measurements of respiration to the stand-level and calculate annual respiratory fluxes. Previous studies have indicated that failure to consider temperature gradients between sun-exposed stems and branches in the crown and shaded lower stems may result in errors when deriving stand-level estimates of stem CO(2) efflux. We measured vertical gradients in sapwood temperature in a mature lowland podocarp rain forest in New Zealand to: (1) estimate the effects of within-stem temperature variation on the vertical distribution of stem CO(2) efflux; and (2) use these findings to estimate stand-level stem CO(2) efflux for this forest. Large within-stem gradients in sapwood temperature (1.6 +/- 0.1 to 6.0 +/- 0.5 degrees C) were observed. However, these gradients did not significantly influence the stand-level estimate of stem CO(2) efflux in this forest (536 +/- 42 mol CO(2) ha(-1) day(-1)) or the vertical distribution of stem CO(2) efflux, because of the opposing effects of daytime warming and nighttime cooling on CO(2) efflux in the canopy, and the small fraction of the woody biomass in the crowns of forest trees. Our findings suggest that detailed measurements of within-stand temperature gradients are unlikely to greatly improve the accuracy of tree- or stand-level estimates of stem CO(2) efflux.     

Tree water storage and its diurnal dynamics related to sap flow and changes in stem volume in old-growth Douglas-fir trees

Diurnal and seasonal tree water storage was studied in three large Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) trees at the Wind River Canopy Crane Research site. Changes in water storage were based on measurements of sap flow and changes in stem volume and tissue water content at different heights in the stem and branches. We measured sap flow by two variants of the heat balance method (with internal heating in stems and external heating in branches), stem volume with electronic dendrometers, and tissue water content gravimetrically. Water storage was calculated from the differences in diurnal courses of sap flow at different heights and their integration. Old-growth Douglas-fir trees contained large amounts of free water: stem sapwood was the most important storage site, followed by stem phloem, branch sapwood, branch phloem and needles. There were significant time shifts (minutes to hours) between sap flow measured at different positions within the transport system (i.e., stem base to shoot tip), suggesting a highly elastic transport system. On selected fine days between late July and early October, when daily transpiration ranged from 150 to 300 liters, the quantity of stored water used daily ranged from 25 to 55 liters, i.e., about 20% of daily total sap flow. The greatest amount of this stored water came from the lower stem; however, proportionally more water was removed from the upper parts of the tree relative to their water storage capacity. In addition to lags in sap flow from one point in the hydrolic pathway to another, the withdrawal and replacement of stored water was reflected in changes in stem volume. When point-to-point lags in sap flow (minutes to hours near the top and stem base, respectively) were considered, there was a strong linear relationship between stem volume changes and transpiration. Volume changes of the whole tree were small (equivalent to 14% of the total daily use of stored water) indicating that most stored water came from the stem and from its inelastic (sapwood) tissues. Whole tree transpiration can be maintained with stored water for about a week, but it can be maintained with stored water from the upper crown alone for no more than a few hours.     

Effects of temperature and tissue nitrogen on dormant season stem and branch maintenance respiration in a young loblolly pine (Pinus taeda) plantation

We measured dormant season (November through February) maintenance respiration rates (R(m)) in stems and branches of 9-year-old loblolly pine (Pinus taeda L.) growing in plots under conditions of controlled nutrient and water supply in an effort to determine the relationships between R(m) and tissue size (surface area, sapwood volume, sapwood dry weight), tissue nitrogen content and temperature. Dormant season R(m) per unit size (i.e., surface area, &mgr;mol m(-2) s(-1); sapwood volume, &mgr;mol m(-3) s(-1); or sapwood dry weight, nmol g(-1) s(-1)) varied with tissue size, but was constant with respect to tissue nitrogen content (&mgr;mol mol(-1) N s(-1)). Cambium temperature accounted for 61 and 77% of the variation in stem and branch respiration, respectively. The basal respiration rate (respiration at 0 degrees C) increased with tissue nitrogen content, however, the Q(10) did not. Improved nutrition more than doubled stem basal respiration rate and increased branch basal respiration by 38%. Exponential equations were developed to model stem and branch respiration as a function of cambium temperature and tissue nitrogen content. We conclude that failure to account for tissue nitrogen effects on respiration rates will result in serious errors when estimating annual maintenance costs.     

Oak decline: I. Fungi associated with various disease symptoms on overground portions of middle-aged and old oak (Quercus robur L.)

Fungi colonizing overground portions of Quercus robur art presented separately for each of the following types of disease symptoms: leaves with atrophy and necrotic areas, dead branches, dead tops of branches, dead water sprouts, local necrotic areas in bark and phloem, locally dead and discoloured sapwood in debarked places, local discolorations inside sapwood, extensive nec-rotic areas on stems, dead trees. Over 80 species of fungi were found in all, mainly from As-comycotina and Deuteromycotina. Connected with each type of symptom there were several species of fungi which were more frequent than the remaining ones. In most cases fungi seem to be of secondary importance in development of symptoms characteristic for oak decline.     

Climate influences the leaf area/sapwood area ratio in Scots pine

We tested the hypothesis that the leaf area/sapwood area ratio in Scots pine (Pinus sylvestris L.) is influenced by site differences in water vapor pressure deficit of the air (D). Two stands of the same provenance were selected, one in western Scotland and one in eastern England, so that effects resulting from age, genetic variability, density and fertility were minimized. Compared with the Scots pine trees at the cooler and wetter site in Scotland, the trees at the warmer and drier site in England produced less leaf area per unit of conducting sapwood area both at a stem height of 1.3 m and at the base of the live crown, whereas stem permeability was similar at both sites. Also, trees at the drier site had less leaf area per unit branch cross-sectional area at the branch base than trees at the wetter site. For each site, the average values for leaf area, sapwood area and permeability were used, together with values of transpiration rates at different D, to calculate average stem water potential gradients. Changes in the leaf area/sapwood area ratio acted to maintain a similar water potential gradient in the stems of trees at both sites despite climatic differences between the sites.     

纵坑切梢小蠹蓝污真菌侵害云南松树组织解剖观察

通过对云南松树进行纵坑切梢小蠹（Tomicus piniperda）蓝污真菌（Leptogramphium yunnanense）接种试验，对云南松受害组织进行了形态解剖观察，验证了蓝污真菌对云南松的致病作用。蓝污真菌发现于受害木质部和韧皮组织中的各类细胞中，在木质部，蓝污区域呈现出以心材为中心的扇形分布。在韧皮组织内，蓝污真菌主要沿着细胞轴向在垂直方向上生长，并可通过孔纹进入到水平方向上的相邻细胞内。本项研究揭示了该蓝污真菌在云南松树干韧皮和木质部组织，细胞内的生长和蔓延规律，进一步表明纵坑切梢小蠹蓝污真菌L.yunnanense是云南松的一种病原真菌。     

Respiratory potential in sapwood of old versus young ponderosa pine trees in the Pacific Northwest

Our primary objective was to present and test a new technique for in vitro estimation of respiration of cores taken from old trees to determine respiratory trends in sapwood. Our secondary objective was to quantify effects of tree age and stem position on respiratory potential (rate of CO2 production of woody tissue under standardized laboratory conditions). We extracted cores from one to four vertical positions in boles of +200-, +50- and +15-year-old Pinus ponderosa Dougl. ex Laws. trees. Cores were divided into five segments corresponding to radial depths of inner bark; outer, middle and inner sapwood; and heartwood. Data suggested that core segment CO2 production was an indicator of its respiratory activity, and that potential artifacts caused by wounding and extraction were minimal. On a dry mass basis, respiratory potential of inner bark was 3-15 times greater than that of sapwood at all heights for all ages (P < 0.0001). Within sapwood at all heights and in all ages of trees, outer sapwood had a 30-60% higher respiratory potential than middle or inner sapwood (P < 0.005). Heartwood had only 2-10% of the respiratory potential of outer sapwood. For all ages of trees, sapwood rings produced in the same calendar year released over 50% more CO2 at treetops than at bases (P < 0.0001). When scaled to the whole-tree level on a sapwood volume basis, sapwood of younger trees had higher respiratory potential than sapwood of older trees. In contrast, the trend was reversed when using the outer-bark surface area of stems as a basis for comparing respiratory potential. The differences observed in respiratory potential calculated on a core dry mass, sapwood volume, or outer-bark surface area basis clearly demonstrate that the resulting trends within and among trees are determined by the way in which the data are expressed. Although these data are based on core segments rather than in vivo measurements, we conclude that the relative differences are probably valid even if the absolute differences are not.     

Neutral lipids and phospholipids in Scots pine (Pinus sylvestris) sapwood and heartwood

Variations in the concentration and composition of triacylglycerols, free fatty acids and phospholipids were analyzed in Scots pine (Pinus sylvestris L.) trees at five sites. Disks were taken at breast height or at a height of 4 m from the stems of 81 trees differing in diameter and growth rate. The mean concentration of triacylglycerols in sapwood was 26 mg g(-1) dry mass; however, variation among trees was large (16-51 mg g(dm)(-1)). The concentration of triacylglycerols was slightly larger at 4 m height in the stem than at breast height. Concentrations of triacylglycerols did not differ between the sapwood of young and small-diameter stems (DBH < 12 cm) and the sapwood of old stems (DBH > 36 cm). Concentrations of free fatty acids were negligible in the outer sapwood, but ranged between 5 and 18 mg g(dm)(-1) in the heartwood. The most abundant fatty acids of triacylglycerols were oleic (18:1), linoleic (18:2omega6, 18:2Delta5,9), linolenic (pinolenic, 18:3Delta5,9,12 and 18:3omega3) and eicosatrienoic acid (20:3Delta5,11,14 and 20:3omega6). The concentration of linoleic acid comprised 39-46% of the triacylglycerol fatty acids and the concentration was higher in the slow-growing stem from northern Finland than in the stems from southern Finland. Major phospholipids were detected only in sapwood, and only traces of lipid phosphorus were detected in heartwood.     

Picea abies sapwood width: Variations within and between trees

Abstract

This study focused on the amount of sapwood and its variation by means of computed tomographic (CT) imaging. Twenty-four trees were selected from four Norway spruce [Picea abies (L.) Karst.] stands in north-eastern France, varying in age, density and fertility. In each stand, sampled trees represented the dominant, co-dominant and suppressed strata. The heartwood/sapwood boundary was detected from the CT images, and the heartwood and sapwood amount and their variations were then evaluated. At the within-tree level sapwood width was relatively constant along the tree stem above the butt swelling and below the living crown. The between-tree sapwood width variations were partially explained by the total cross-sectional area of living branches. This result opens up the possibility of investigating within-tree allometric relationships. Sapwood width was found to be highly correlated with tree slendemess (tree height/breast height diameter) and with the relative height of the crown. This suggests that sapwood width could be readily predicted from conventional forest inventory measurements. The number of sapwood rings within the stem was largely dependent on cambial age, and could be determined dynamically using the concept of mean lifetime of sapwood rings.     

Extending sapwood — Leaf area relationships from stems to roots in Coast Douglas-fir

? Studies of allometric relationships between leaf area and the cross-sectional area (CSA) of sapwood in the stem have shed light on the structural and functional relationships between water-conducting and photosynthetic tissues.

? The purpose of this study was to test whether sapwood-leaf area relationships could be extended from stems to roots in coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Twelve trees were felled, their stumps were excavated, and the CSA of sapwood and heartwood were estimated for individual roots, entire root systems, and stem section.

? Root sapwood CSA was greater than sapwood CSA throughout the stem, and the ratio of leaf area to sapwood CSA (A _l :A _s ) was accordingly lower for root sapwood. The relationship between sapwood CSA and leaf area was more variable in roots and at groundline compared to crown base. Root A _l :A _s decreased with relative tree height (tree height/mean stand height).

? The strong allometric relationship between leaf area and the CSA of sapwood in the stem generally holds when extended to roots. The greater CSA of sapwood in roots versus stems may reflect differences in their roles in supporting the tree.

     

Response of LAI-2000 estimates to changes in plant surface area index in a Scots pine stand

We assessed the accuracy with which the LAI-2000 plant canopy analyzer measured changes in leaf area index (LAI) and plant area index (PAI) in a 25-year-old Scots pine (Pinus sylvestris L.) stand. Stand density was 2100 stems ha(-1) and mean tree height was 8.7 m. Needle and branch areas of the stand were reduced progressively to zero by the stepwise removal of branches on all trees growing in a circular plot with a radius of 25 m. An LAI-2000 estimate was taken after each step reduction. The needle and branch surface areas removed at each step were estimated from direct measurements and were compared with the changes in the LAI-2000 estimates. Initially (before removal of branches), directly measured PAI was 5.2 (needles = 86%, branches = 8% and stems = 6%). The LAI-2000 estimate of total surface area was 66% of direct PAI and 77% of direct LAI. There was a nonlinear relationship between the LAI-2000 estimate and directly measured PAI, such that their ratio (equivalent to the clumping factor) increased from 0.66 to 1.05 with decreasing PAI. At the last measurement, when only stems were left, the LAI-2000 estimate agreed well with the direct measurement of PAI. The LAI-2000 underestimated the direct measurement of LAI at the first three steps when LAI was > 2 and the proportion of woody area was small (< 20%). However, because the LAI-2000 estimate included stem and branch areas, it overestimated the direct measurement of LAI at the last three measurements when the proportion of woody area was large (> 20%).     

A method for reconstructing the development of the sapwood area of balsam fir

Leaf area is commonly estimated as a function of sapwood area. However, because sapwood changes to heartwood over time, it has not previously been possible to reconstruct either the sapwood area or the leaf area of older trees into the past. In this study, we report a method for reconstructing the development of the sapwood area of dominant and codominant balsam fir (Abies balsamea (L.) Mill.). The technique is based on establishing a species-specific relationship between the number of annual growth rings in the sapwood area and tree age. Because the number of annual growth rings in the sapwood of balsam fir at a given age was found to be independent of site quality and stand density, the number of rings in sapwood (NRS) can be predicted from the age of a tree thus: NRS = 14.818 (1 - e(-0.031 age)), unweighted R(2) = 0.80, and NRS = 2.490 (1 - e(-0.038 age)), unweighted R(2) = 0.64, for measurements at breast height and at the base of the live crown, respectively. These nonlinear asymptotic regression models based only on age, were not improved by adding other tree variables such as diameter at breast height, diameter at the base of the live crown, total tree height or percent live crown.     

Killing effect and translocation of picloram in the stem of aspen,Populus tremula

Picloram was injected to different depths in the stems of aspen trees during spring and summer, and the killing effect on the foliage was studied. The efficacy of the applied dosage was correlated better to the surface area and volume of the stem than to the diameter. The young foliage was affected first by the herbicide. Both the xylem and the phloem were involved in the effects that resulted from translocation of the herbicide.     

Internal remobilization of carbohydrates, lipids, nitrogen and phosphorus in the Mediterranean evergreen oak Quercus ilex

Remobilization of internal resources is an important mechanism enabling plants to be partly independent of external nutrient availability. We assessed resource remobilization during the growing period in woody and foliar tissues of leafy branches of mature evergreen Mediterranean oak (Quercus ilex L.) at three field sites. We compared nonstructural carbohydrates, lipids, nitrogen and phosphorus pools in leaves and stems before bud burst (March) and at the end of the growing period (July). We also experimentally defoliated leafy branches to determine the storage function of old leaves. Changes in pools of carbon compounds in leaves and stems during spring and in response to defoliation indicated that foliar and woody tissues could provide carbon to support shoot growth. Independently of stem age, soluble sugar and lipid pools decreased significantly during spring. Changes in leaf pools between March and July involved all compounds measured except starch and were accompanied by a 5% decrease in mean leaf biomass. During the same period, 15% of the nitrogen and 25% of the phosphorus were removed from leaves. In contrast, woody tissues did not remobilize nitrogen or phosphorus. Our results support earlier hypotheses that leaves of evergreen species have a primary role in resource remobilization.     

Systematic variation in xylem hydraulic capacity within the crown of white ash (Fraxinus americana)

A 7-m tall white ash tree (Fraxinus americana Marsh.) was dissected, and hydraulic parameters of the xylem were determined by inducing a steady-state flow of water through the stem segments and monitoring volume and velocity flow rates. Leaf-specific conductivity (LSC) was highest in the main stem and lowest in some of the leaf-bearing lateral shoots. The LSC was higher in the main stem than in branches and higher in primary than in secondary branch axes. Terminal leaf-bearing shoots were larger and had a significantly greater mean LSC than subjacent lateral shoots. A significant reduction in LSC was associated with the transition between 1- and 2-year-old growth. In branches of the same age, there was a close correspondence among LSC, branch position and branch size. The average LSC of leaf-bearing shoots from south-facing branches was 43% greater than that of shoots from north-facing branches. Within-crown variation in LSC was associated with variation in velocity flow rate (V). By contrast, the ratio of potentially functional xylem area to supported leaf area (A(pf)/A(l)) was relatively stable throughout the crown. Stratification of stems by Strahler order accounted for approximately 70% of the total variation in LSC. These results suggest that (1) there exists a systematic pattern of variation in LSC distribution within the crown of white ash, (2) within-crown variability in LSC is primarily the result of variation in mean vessel diameter, and (3) there is a physiological linkage between LSC and crown morphology that is maintained through a positive feedback mechanism during branch ontogeny.     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Correlations between stable carbon-isotope abundance and hydraulic conductivity in Douglas-fir across a climate gradient in Oregon, USA

Stomatal conductance in trees is related to both foliar carbon-isotope abundance and stem hydraulic properties. By combining these relationships, I hypothesized that carbon-isotope abundance in foliage should vary with limitations to water movement through supporting branches. I sampled Douglas-fir branches (Pseudotsuga menziesii (Mirb.) Franco) from six sites across a climate gradient in Oregon, USA for foliar carbon-isotope abundance and stem hydraulic properties. I used a forest growth model to quantify climate-induced stomatal limitations, expressed as reduced potential transpiration, across the gradient. Foliar stable carbon-isotope abundance showed a strong inverse relationship with branch specific conductivity (hydraulic conductivity per unit functional sapwood area) and leaf-specific conductivity (hydraulic conductivity per unit leaf area). Foliar stable carbon-isotope abundance was correlated with modeled reductions in potential transpiration; however, the inclusion of leaf-specific conductivity improved the correlation by more than 30%. Combined, leaf-specific conductivity and climate-induced stomatal constraints explained 84% of the variation in foliar isotope abundance in 1994 foliage. This model was confirmed on foliage classes 1990-1993.