A mathematical model linking tree sap flow dynamics to daily stem diameter fluctuations and radial stem growth

To date, models for simulating sap flow dynamics in individual trees with a direct link to stem diameter variation include only the diameter fluctuation driven by a change in stem water storage. This paper reports results obtained with a comprehensive flow and storage model using whole-tree leaf transpiration as the only input variable. The model includes radial stem growth based on Lockhart's equation for irreversible cell expansion. It was demonstrated that including growth is essential to obtaining good simulation results. To model sap flow dynamics, capacitance of storage tissues was assumed either constant (i.e., electrical analogue approach) or variable and dependent on the water content of the respective storage tissue (i.e., hydraulic system approach). These approaches resulted in different shapes for the desorption curve used to calculate the capacitance of storage tissues. Comparison of these methods allowed detection of specific differences in model simulation of sap flow at the stem base (F(stem)) and stem diameter variation (D). Sensitivity analysis was performed to select a limited subset of identifiable parameters driving most of the variability in model predictions of F(stem) and D Both the electrical analogue and the hydraulic system approach for the flow and storage model were successfully calibrated and validated for the case of a young beech tree (Fagus sylvatica L.). Use of an objective model selection criterion revealed that the flow and storage model based on the electrical analogue approach yielded better predictions.     

Nocturnal sap flow characteristics and stem water recharge of Acacia mangium

In this paper, we studied the nocturnal stem water recharge of Acacia mangium. It is helpful to improve the precision of canopy transpiration estimation and canopy stomatal conductance, and to further understand the lag time of canopy transpiration to stem sap flow. In this study, the whole-tree sap flow in an A. mangium forest was measured by using Granier’s thermal dissipation probe for over two years in the hilly land of South China. The environmental factors, including relative humidity (RH), precipitation, vapor pressure deficit (VPD), photosynthetically active radiation (PAR), and air temperature (T _a) were recorded simultaneously. The stem water recharge of A. mangium was analyzed on both daily and monthly scales. Sap flux density was lower at night than during the day. The time range of nighttime sap flux density was longer in the dry season than in the wet season. The water recharging mainly occurred from sunset to midnight. No significant differences were observed among inter-annual nighttime water recharges. Nighttime water recharge had no significant correlation with environmental factors, but was well correlated with the diameter at breast height, tree height, and crown size. In the dry season the contribution of nighttime water recharge to total transpiration had significant correlations with daytime transpiration, total transpiration, VPD, PAR and T _a, while in the wet season it was significantly correlated with daily transpiration and total transpiration. __________ Translated from Chinese Journal of Ecology, 2007, 26(4): 476–482 [译自: 生态学杂志]     

Link between diurnal stem radius changes and tree water relations

Internal water reserves are depleted and replenished daily, not only in succulent plants, but also in trees. The significance of these changes in tissue water storage for tree water relations was investigated by monitoring diurnal fluctuations in stem radius. In 6-year-old potted Norway spruce (Picea abies (L.) Karst.) trees, whole-tree transpiration rate (T), sap flow at the stem base and fluctuations in stem radius were measured at 10-min intervals over eight successive weeks. The dynamics of diurnal water storage in relation to the daily course of water movement was simulated and the contribution of stored water to T quantified. The finding that, in P. abies, the course of bark water content is linearly coupled to stem radius fluctuations provided the basis for linking stem radius changes to a functional flow and storage model for tree water relations. This model, which consists of physical functions only and is driven by a single input variable (T), accurately simulates the diurnal course of changes in stem radius and water storage of the tree crown and stem. It was concluded that fluctuations were mainly determined by the course of transpiration. The availability of soil water and the degree to which storage tissues were saturated were also factors affecting the diurnal course of stem radius changes. Internally stored water contributed to daily transpiration even in well-watered trees, indicating that stored water plays an important role not only during periods of drought, but whenever water transport occurs within the tree. Needle and bark water reserves were most heavily depleted during transpiration. Together they supplied approximately 10% of daily T on sunny days, and up to 65% on cloudy days. On a daily basis, the crown (mainly needles) contributed approximately eight times more water to T than the stem (mainly bark). The depletion of the two storage pools and the water movements observed in the trees always occurred in the same sequence. In the morning, T first caused a depletion of the water stored in the crown. It then caused depletion of bark storage tissues at ever increasing distances from the needles. Up to 75% of the transpired water could be withdrawn from storage tissues when the increase in T reached a maximum.     

Diurnal and seasonal variability in the radial distribution of sap flow: predicting total stem flow in Pinus taeda trees

We monitored the radial distribution of sap flux density (v; g H2O m(-2) s(-1)) in the sapwood of six plantation-grown Pinus taeda L. trees during wet and dry soil periods. Mean basal diameter of the 32-year-old trees was 33.3 cm. For all trees, the radial distribution of sap flow in the base of the stem (i.e., radial profile) was Gaussian in shape. Sap flow occurred maximally in the outer 4 cm of sapwood, comprising 50-60% of total stem flow (F), and decreased toward the center, with the innermost 4 cm of sapwood (11-15 cm) comprising less than 10% of F. The percent of flow occurring in the outer 4 cm of sapwood was stable with time (average CV < 10%); however, the percentage of flow occurring in the remaining sapwood was more variable over time (average CV > 40%). Diurnally, the radial profile changed predictably with time and with total stem flow. Seasonally, the radial profile became less steep as the soil water content (theta) declined from 0.38 to 0.21. Throughout the season, daytime sap flow also decreased as theta decreased; however, nighttime sap flow (an estimate of stored water use) remained relatively constant. As a result, the percentage of stored water use increased as theta declined. Time series analysis of 15-min values of F, theta, photosynthetically active radiation (PAR) and vapor pressure deficit (D) showed that F lagged behind D by 0-15 min and behind PAR by 15-30 min. Diurnally, the relationship between F and D was much stronger than the relationship between F and PAR, whereas no relationship was found between F and theta. An autoregressive moving average (ARIMA) model estimated that 97% of the variability in F could be predicted by D alone. Although total sap flow in all trees responded similarly to D, we show that the radial distribution of sap flow comprising total flow could change temporally, both on daily and seasonal scales.     

Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees

In large trees, the daily onset of transpiration causes water to be withdrawn from internal storage compartments, resulting in lags between changes in transpiration and sap flow at the base of the tree. We measured time courses of sap flow, hydraulic resistance, plant water potential and stomatal resistance in co-occurring tropical forest canopy trees with trunk diameters ranging from 0.34-0.98 m, to determine how total daily water use and daily reliance on stored water scaled with size. We also examined the effects of scale and tree hydraulic properties on apparent time constants for changes in transpiration and water flow in response to fluctuating environmental variables. Time constants for water movement were estimated from whole-tree hydraulic resistance (R) and capacitance (C) using an electric circuit analogy, and from rates of change in water movement through intact trees. Total daily water use and reliance on stored water were strongly correlated with trunk diameter, independent of species. Although total daily withdrawal of water from internal storage increased with tree size, its relative contribution to the daily water budget (approximately 10%) remained constant. Net withdrawal of water from storage ceased when upper branch water potential corresponded to the sapwood water potential (Psi(sw)) at which further withdrawal of water from sapwood would have caused Psi(sw) to decline precipitously. Stomatal coordination of vapor and liquid phase resistances played a key role in limiting stored water use to a nearly constant fraction of total daily water use. Time constants for changes in transpiration, estimated as the product of whole- tree R and C, were similar among individuals (~0.53 h), indicating that R and C co-varied with tree size in an inverse manner. Similarly, time constants estimated from rates of change in crown and basal sap flux were nearly identical among individuals and therefore independent of tree size and species.     

雪花梨树干液流特征及其与环境因子的关系

采用热扩散茎流计法对雪花梨的树干液流进行了研究。结果表明,雪花梨树干边材液流曲线呈宽峰状,表现出明显的昼夜变化和季节变化规律,其变化规律和气象因子关系密切,特别是大气温度、空气湿度和辐射强度,土壤水分则主要从耗水总量角度去影响树干液流。     

Tree stem diameter variations and transpiration in Scots pine: an analysis using a dynamic sap flow model

A dynamic model for simulating water flow in a Scots pine (Pinus sylvestris L.) tree was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration, together with the elasticity of wood tissue, causes variations in the diameter of a tree stem and branches. The change in xylem diameter can be linked to water tension in accordance with Hookea s law. The model was tested against field measurements of the diurnal xylem diameter change at different heights in a 37-year-old Scots pine at Hyyti?l?, southern Finland (61 degrees 51' N, 24 degrees 17' E, 181 m a.s.l.). Shoot transpiration and soil water potential were input data for the model. The biomechanical and hydraulic properties of wood and fine root hydraulic conductance were estimated from simulated and measured stem diameter changes during the course of 1 day. The estimated parameters attained values similar to literature values. The ratios of estimated parameters to literature values ranged from 0.5 to 0.9. The model predictions (stem diameters at several heights) were in close agreement with the measurements for a period of 6 days. The time lag between changes in transpiration rate and in sap flow rate at the base of the tree was about half an hour. The analysis showed that 40% of the resistance between the soil and the top of the tree was located in the rhizosphere. Modeling the water tension gradient and consequent woody diameter changes offer a convenient means of studying the link between wood hydraulic conductivity and control of transpiration.     

Estimating stand water use of large mountain ash trees and validation of the sap flow measurement technique

Mountain ash (Eucalyptus regnans F.J. Muell.) forest catchments exhibit a strong relationship between stand age and runoff, attributed inter alia to differences in tree water use. However, the tree water use component of the mountain ash forest water balance is poorly quantified. We have used the sap flow technique to obtain estimates of daily water use in large mountain ash trees. First, the sap flow technique was validated by means of an in situ cut tree experiment. Close agreement was obtained between the sap flow estimate of water use and the actual uptake of water by the tree from a reservoir. Second, we compared the variability in sap velocity between a symmetric and an asymmetric tree by using multiple sap flow loggers. In the symmetric tree, velocity was fairly uniform throughout the xylem during the day, indicating that accurate sap flow estimates can be obtained with a minimal number of sampling points. However, large variations in sap velocity were observed in the asymmetric tree, indicating that much larger sampling sizes are required in asymmetric stems for an accurate determination of mean sap velocity. Finally, we compared two procedures for scaling individual tree sap flow estimates to the stand level based on stem diameter and leaf area index measurements. The first procedure was based on a regression between stem diameter and tree water use, developed on a small sample of trees and applied to a stand-level census of stem diameter values. Inputs to the second procedure were tree water use and leaf area of a single tree and the leaf area index of the stand. The two procedures yielded similar results; however, the first procedure was more robust but it required more sampling effort than the second procedure.     

Daily shoot extension growth of peach trees growing on rootstocks that reduce scion growth is related to daily dynamics of stem water potential

We studied relationships between diurnal patterns of stem water potential (PsiSTEM) and stem extension growth of the same scion cultivar growing on three rootstocks with differing size-controlling potentials. The peach trees (Prunus persica (L.) Batsch) used in this field experiment consisted of an early-maturing freestone cultivar, 'Flavorcrest,' grafted onto three different rootstocks: Nemaguard (a vigorous seed-propagated control, P. persica x P. davidiana hybrid), Hiawatha (an intermediate vigor rootstock, derived from an open pollinated seedling of a P. besseyi x P. salicina hybrid) and K-146-43 (a semi-dwarfing rootstock, P. salicina x P. persica hybrid). Diurnal patterns of PsiSTEM and stem extension growth were measured on six dates (March 29, April 12, April 26, May 10, May 24 and June 18) during the primary period of peach shoot extension growth. Rootstocks clearly affected diurnal patterns of PsiSTEM and stem extension growth. Trees on K-146-43 had the lowest midday PsiSTEM and stem extension growth. Differences among rootstocks in the amount of diurnal oscillation in PsiSTEM explained stem extension rate differences induced by the three rootstocks. The sensitivity of shoot extension growth to tree water relations tended to decrease as the season progressed and was not apparent by mid-June. The results of the study indicate that water relations may play an important role in the dwarfing mechanism induced by size-controlling peach rootstocks.     

Traits related to stem volume yield capacity in sample trees of pinus sylvestris L. and pinus contorta dougl.

Species, provenances and even individual trees are known to differ in stem volume yield capacity, even when grown on the same site. Increased knowledge of the fundamental mechanisms for different productivity is essential for the success of breeding research and for the maximization of yield. Genetically homogeneous tree materials were selected on the basis of expected differences in stem volume yield capacity and trees were sampled and destructively analysed. The hypothesis to be tested was that yield capacity is correlated to certain traits of trees. High correlations with stem volume yield capacity were shown for harvest index, needle‐use efficiency and mineral‐nutrient‐use efficiency. Only weak correlations were shown for needle longevity and nitrogen concentration in needles. Also, indications were found that phenotypic plasticity was correlated positively to stem volume growth capacity. An attempt to correlate the ability to retranslocate nitrogen from dying needles with yield capacity failed.     

Diurnal and seasonal changes in stem increment and water use by yellow poplar trees in response to environmental stress

To evaluate indicators of whole-tree physiological responses to climate stress, we determined seasonal, daily and diurnal patterns of growth and water use in 10 yellow poplar (Liriodendron tulipifera L.) trees in a stand recently released from competition. Precise measurements of stem increment and sap flow made with automated electronic dendrometers and thermal dissipation probes, respectively, indicated close temporal linkages between water use and patterns of stem shrinkage and swelling during daily cycles of water depletion and recharge of extensible outer-stem tissues. These cycles also determined net daily basal area increment. Multivariate regression models based on a 123-day data series showed that daily diameter increments were related negatively to vapor pressure deficit (VPD), but positively to precipitation and temperature. The same model form with slight changes in coefficients yielded coefficients of determination of about 0.62 (0.57-0.66) across data subsets that included widely variable growth rates and VPDs. Model R2 was improved to 0.75 by using 3-day running mean daily growth data. Rapid recovery of stem diameter growth following short-term, diurnal reductions in VPD indicated that water stored in extensible stem tissues was part of a fast recharge system that limited hydration changes in the cambial zone during periods of water stress. There were substantial differences in the seasonal dynamics of growth among individual trees, and analyses indicated that faster-growing trees were more positively affected by precipitation, solar irradiance and temperature and more negatively affected by high VPD than slower-growing trees. There were no negative effects of ozone on daily growth rates in a year of low ozone concentrations.     

Impact of soil drought on sap flow and water status of evergreen trees in a tropical monsoon forest in northern Thailand

Hill evergreen forest is the dominant vegetation type in northern Thailand. In this region, there is higher atmospheric evaporative demand and lower soil moisture during the 5- to 7-month dry season than in the rainy season under influences from Asian monsoons. In an earlier study we revealed that canopy-scale transpiration is actively maintained even during the latter part of the dry season in hill evergreen forest. However, the impact of soil drought on tree water use was not investigated. To clarify the ecohydrological processes at this site, we used individual tree-scale measurements during a 2-year period to base our examination of whether limited water use in individual trees is caused by soil drought in the latter part of the dry season. Sap flow and water potential measurements were conducted in four evergreen trees, two large emergent trees 29.8 and 25.4 m high, and two smaller understory trees 4.8 and 1.4 m high.The amount of rainfall preceding the late dry season of 2004 was significantly less than that preceding the late dry season of 2003. Although a distinct decrease in sap-flow velocities in individual trees due to soil water stress was not found in the late dry season of 2003, it did become comparatively apparent in the late dry season of 2004; ranging from 10 to 40% for a given atmospheric evaporative demand. Furthermore, the reductions in sap-flow velocities and predawn stem-water potential were most significant in the smallest tree. The recovery of sap-flow velocities and water potential in the smallest tree after irrigation confirmed that the reductions in sap-flow velocity and predawn stem-water potential in the smallest tree were caused by soil drought. These results suggest that shallower roots could be reason for the significant decrease in water use in the smallest trees. The deeper roots of larger trees could be the reason for the reduced impact of soil drought on water use in larger trees, and canopy-scale transpiration might be maintained by larger trees, even in an unusually severe drought. These possibilities provide a new insight for management of evergreen forests under Asian monsoon influences.     

Critical effects on the photosynthetic efficiency and stem sap flow of poplar in the Yellow River Delta in response to soil water

To explore the critical relationships of photo-synthetic efficiency and stem sap flow to soil moisture,two-year-old poplar saplings were selected and a packaged...     

Spatial sap flow and xylem anatomical characteristics in olive trees under different irrigation regimes

The compensation heat pulse (CHP) method is widely used to estimate sap flow and transpiration in conducting organs of woody plants. Previous studies have reported a natural azimuthal variability in sap flow, which could have practical implications in locating the CHP probes and integrating their output. Sap flow of several olive trees (Olea europaea L. cv. 'Arbequina') previously grown under different irrigation treatments were monitored by the CHP method, and their xylem anatomical characteristics were analyzed from wood samples taken at the same location in which the probes were installed. A significant azimuthal variability in the sap flow was found in a well-irrigated olive tree monitored by eight CHP probes. The azimuthal variability was well related to crown architecture, but poorly to azimuthal differences in the xylem anatomical characteristics. Well-irrigated and deficit-irrigated olive trees showed similar xylem anatomical characteristics, but they differed in xylem growth and in the ratio of nocturnal-to-diurnal sap flow (N/D index). The results of this work indicate that transpiration cannot be accurately estimated by the CHP method in olive trees if a small number of sensors are employed and that the N/D index could be used as a sensitive water status indicator.     

Variability with xylem depth in sap flow in trunks and branches of mature olive trees

Knowledge of sap flow variability in tree trunks is important for up-scaling transpiration from the measuring point to the whole-tree and stand levels. Natural variability in sap flow, both radial and circumferential, was studied in the trunks and branches of mature olive trees (Olea europea L., cv Coratina) by the heat field deformation method using multi-point sensors. Sapwood depth ranged from 22 to 55 mm with greater variability in trunks than in branches. Two asymmetric types of sap flow radial patterns were observed: Type 1, rising to a maximum near the mid-point of the sapwood; and Type 2, falling continuously from a maximum just below cambium to zero at the inner boundary of the sapwood. The Type 1 pattern was recorded more often in branches and smaller trees. Both types of sap flow radial patterns were observed in trunks of the sample trees. Sap flow radial patterns were rather stable during the day, but varied with soil water changes. A decrease in sap flow in the outermost xylem was related to water depletion in the topsoil. We hypothesized that the variations in sap flow radial pattern in a tree trunk reflects a vertical distribution of water uptake that varies with water availability in different soil layers.     

Temporal dynamics of stem expansion and contraction in savanna trees: withdrawal and recharge of stored water

Relationships between diel changes in stem expansion and contraction and discharge and refilling of stem water storage tissues were studied in six dominant Neotropical savanna (cerrado) tree species from central Brazil. Two stem tissues were studied, the active xylem or sapwood and the living tissues located between the cambium and the cork, made up predominantly of parenchyma cells (outer parenchyma). Outer parenchyma and sapwood density ranged from 320 to 410 kg m(-3) and from 420 to 620 kg m(-3), respectively, depending on the species. The denser sapwood tissues exhibited smaller relative changes in cross-sectional area per unit change in water potential compared with the outer parenchyma. Despite undergoing smaller relative changes in cross-sectional area, the sapwood released about 3.5 times as much stored water for a given change in area as the outer parenchyma. Cross-sectional area decreased earlier in the morning in the outer parenchyma than in the sapwood with lag times up to 30 min for most species. The relatively small lag time between dimensional changes of the two tissues suggested that they were hydraulically well connected. The initial morning increase in basal sap flow lagged about 10 to 130 min behind that of branch sap flow. Species-specific lag times between morning declines in branch and main stem cross-sectional area were a function of relative stem water storage capacity, which ranged from 16 to 31% of total diurnal water loss. Reliance on stored water to temporarily replace transpirational losses is one of the homeostatic mechanisms that constrain the magnitude of leaf water deficits in cerrado trees.     

Use of water stress integral to evaluate relationships between soil moisture,plant water stress and stand productivity in young Douglas-fir trees

Water stress integral (WSI) is a method of assessing cumulative plant water stress over a chosen period of time. While the technique has been used in other tree species, it has not been applied for reforestation projects. In this study we used the WSI approach for newly planted Douglas-fir in the Pacific Northwest (USA), where the Mediterranean climate, plant community development, and competition for water all play key roles in the success of establishment efforts. In this study, previously reported seedling growth, xylem water potential, and soil moisture data were utilized to provide direct correlations between Douglas-fir productivity, soil water availability and WSI. For each growing season, a strong relationship between WSI and volume growth as well as a strong linear relationship between WSI and soil moisture measured during mid-August was found. On average, for each reduction of 0.01 cm³ cm^?3 in soil moisture measured during mid-August, Douglas-fir seedling volume growth decreased by 5.6 and 7.7% in the first and second growing seasons, respectively. Preserving soil moisture until early-August through the judicial application of vegetation management regimes was critical for maximizing stand productivity. Based on these results, a single evaluation of soil volumetric water content during early-August can be used as a predictor of stand productivity during the initial two seasons of forest establishment.     

Daily sap flow and maximum daily trunk shrinkage measurements for diagnosing water stress in early maturing peach trees during the post-harvest period

We compared the sensitivity of two continuously recorded plant-based water stress indicators (sap flow, SF, and maximum daily trunk shrinkage, MDS) to detect changes in the water status of 4-year-old early maturing peach trees (Prunus persica (L.) Batsch cv. Flordastar grafted on GF-677 peach rootstock) during a cycle of deficit irrigation and recovery. The feasibility of obtaining SF and MDS reference equations for use in irrigation scheduling during the post-harvest period was also studied in trees irrigated in excess of crop water requirements. We found that MDS was a more sensitive and reliable detector of changes in plant water status than SF, making it a more precise tool for irrigation scheduling. Baseline relationships between SF or MDS and the climatic variables (air temperature, vapor pressure deficit (VPD) and crop reference evapotranspiration (ETo)) were established, despite some scatter in the data. Among the climatic variables, SF correlated more closely with ETo, whereas MDS correlated more closely with mean daily air temperature (T (m)). The fits of the regressions between MDS and ETo, midday air temperature and T (m) for individual periods were better than those obtained in the overall regressions, confirming that daily stem diameter variations must be considered not only in the context of plant water status but also in the context of plant carbon status.     

Variation in water potential, hydraulic characteristics and water source use in montane Douglas-fir and lodgepole pine trees in southwestern Alberta and consequences for seasonal changes in photosynthetic capacity

Tree species response to climate change-induced shifts in the hydrological cycle depends on many physiological traits, particularly variation in water relations characteristics. We evaluated differences in shoot water potential, vulnerability of branches to reductions in hydraulic conductivity, and water source use between Pinus contorta Dougl. ex Loud. var. latifolia Engelm. (lodgepole pine) and Pseudotsuga menziesii (Mirb.) Franco (interior Douglas-fir), and determined the consequences for seasonal changes in photosynthetic capacity. The Douglas-fir site had soil with greater depth, finer texture and higher organic matter content than soil at the lodgepole pine site, all factors that increased the storage of soil moisture. While the measured xylem vulnerability curves were quite similar for the two species, Douglas-fir had lower average midday shoot water potentials than did lodgepole pine. This implied that lodgepole pine exhibited stronger stomatal control of transpiration than Douglas-fir, which helped to reduce the magnitude of the water potential gradient required to access water from drying soil. Stable hydrogen isotope measurements indicated that Douglas-fir increased the use of groundwater during mid-summer when precipitation inputs were low, while lodgepole pine did not. There was a greater reduction of photosynthetic carbon gain in lodgepole pine compared with Douglas-fir when the two tree species were exposed to seasonal declines in soil water content. The contrasting patterns of seasonal variation in photosynthetic capacity observed for the two species were a combined result of differences in soil characteristics at the separate sites and the inherent physiological differences between the species.     

Radial variation in sap velocity as a function of stem diameter and sapwood thickness in yellow-poplar trees

Canopy transpiration and forest water use are frequently estimated as the product of sap velocity and cross-sectional sapwood area. Few studies, however, have considered whether radial variation in sap velocity and the proportion of sapwood active in water transport are significant sources of uncertainty in the extrapolation process. Therefore, radial profiles of sap velocity were examined as a function of stem diameter and sapwood thickness for yellow-poplar (Liriodendron tulipifera L.) trees growing on two adjacent watersheds in eastern Tennessee. The compensation heat pulse velocity technique was used to quantify sap velocity at four equal-area depths in 20 trees that ranged in stem diameter from 15 to 69 cm, and in sapwood thickness from 2.1 to 14.8 cm. Sap velocity was highly dependent on the depth of probe insertion into the sapwood. Rates of sap velocity were greatest for probes located in the two outer sapwood annuli (P1 and P2) and lowest for probes in closest proximity to the heartwood (P3 and P4). Relative sap velocities averaged 0.98 at P1, 0.66 at P2, 0.41 at P3 and 0.35 at P4. Tree-specific sap velocities measured at each of the four probe positions, divided by the maximum sap velocity measured (usually at P1 or P2), indicated that the fraction of sapwood functional in water transport (f(S)) varied between 0.49 and 0.96. There was no relationship between f(S) and sapwood thickness, or between f(S) and stem diameter. The fraction of functional sapwood averaged 0.66 +/- 0.13 for trees on which radial profiles were determined. No significant depth-related differences were observed for sapwood density, which averaged 469 kg m(-3) across all four probe positions. There was, however, a significant decline in sapwood water content between the two outer probe positions (1.04 versus 0.89 kg kg(-1)). This difference was not sufficient to account for the observed radial variation in sap velocity. A Monte-Carlo analysis indicated that the standard error in estimated mean f(S) declined rapidly with increasing sample size. At n = 10, the coefficient of variation in mean f(S) was 7% and at n = 15 it was slightly less than 5%. These observations indicate that radial variation in sap velocity is an important, albeit often overlooked, source of uncertainty in the scaling process. Failure to recognize that not all sapwood is functional in water transport will introduce systematic bias into estimates of both tree and stand water use. Future studies should devise sampling strategies for assessing radial variation in sap velocity and such strategies should be used to identify the magnitude of this variation in a range of non-, diffuse- and ring-porous trees.