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.     

Heat dissipation sensors of variable length for the measurement of sap flow in trees with deep sapwood

Robust thermal dissipation sensors of variable length (3 to 30 cm) were developed to overcome limitations to the measurement of radial profiles of sap flow in large-diameter tropical trees with deep sapwood. The effective measuring length of the custom-made sensors was reduced to 1 cm at the tip of a thermally nonconducting shaft, thereby minimizing the influence of nonuniform sap flux density profiles across the sapwood. Sap flow was measured at different depths and circumferential positions in the trunks of four trees at the Parque Natural Metropolitano canopy crane site, Panama City, Republic of Panama. Sap flow was detected to a depth of 24 cm in the trunks of a 1-m-diameter Anacardium excelsum (Bertero & Balb. ex Kunth) Skeels tree and a 0.65-m-diameter Ficus insipida Willd. tree, and to depths of 7 cm in a 0.34-m-diameter Cordia alliodora (Ruiz & Pav.) Cham. trunk, and 17 cm in a 0.47-m-diameter Schefflera morototoni (Aubl.) Maguire, Steyerm. & Frodin trunk. Sap flux density was maximal in the outermost 4 cm of sapwood and declined with increasing sapwood depth. Considerable variation in sap flux density profiles was observed both within and among the trees. In S. morototoni, radial variation in sap flux density was associated with radial variation in wood properties, particularly vessel lumen area and distribution. High variability in radial and circumferential sap flux density resulted in large errors when measurements of sap flow at a single depth, or a single radial profile, were used to estimate whole-plant water use. Diurnal water use ranged from 750 kg H2O day-1 for A. excelsum to 37 kg H2O day-1 for C. alliodora.     

Diurnal and seasonal variability in radial distribution of sap flux density: Implications for estimating stand transpiration

Daily and seasonal patterns in radial distribution of sap flux density were monitored in six trees differing in social position in a mixed coniferous stand dominated by silver fir (Abies alba Miller) and Norway spruce (Picea abies (L.) Karst) in the Alps of northeastern Italy. Radial distribution of sap flux was measured with arrays of 1-cm-long Granier probes. The radial profiles were either Gaussian or decreased monotonically toward the tree center, and seemed to be related to social position and crown distribution of the trees. The ratio between sap flux estimated with the most external sensor and the mean flux, weighted with the corresponding annulus areas, was used as a correction factor (CF) to express diurnal and seasonal radial variation in sap flow. During sunny days, the diurnal radial profile of sap flux changed with time and accumulated photosynthetic active radiation (PAR), with an increasing contribution of sap flux in the inner sapwood during the day. Seasonally, the contribution of sap flux in the inner xylem increased with daily cumulative PAR and the variation of CF was proportional to the tree diameter, ranging from 29% for suppressed trees up to 300% for dominant trees. Two models were developed, relating CF with PAR and tree diameter at breast height (DBH), to correct daily and seasonal estimates of whole-tree and stand sap flow obtained by assuming uniform sap flux density over the sapwood. If the variability in the radial profile of sap flux density was not accounted for, total stand transpiration would be overestimated by 32% during sunny days and 40% for the entire season.     

Integration of sapflow velocity to estimate plant water use

The heat pulse technique provides an estimate of sapflow velocity at one position within the xylem of an individual plant. Previous experience has shown that the velocity profile across the conducting area cannot be assumed to be constant, necessitating several such point estimates for a reasonable characterization of the velocity profile with depth. These point estimates must be integrated over the sapwood conducting area to derive flux. The most common method is to fit a one degree of freedom least-squares polynomial to the point estimates, and then to integrate this equation across the sapwood profile and around the bole. An alternative method is presented based on a weighted average approach. The latter is shown to be a more robust estimator of flux when velocity profiles exhibit large curvatures.     

Radial patterns of sap flow in woody stems of dominant and understory species: scaling errors associated with positioning of sensors

We studied sap flow in dominant coniferous (Pinus sylvestris L.) and broadleaf (Populus canescens L.) species and in understory species (Prunus serotina Ehrh. and Rhododendron ponticum L.) by the heat field deformation (HFD) method. We attempted to identify possible errors arising during flow integration and scaling from single-point measurements to whole trees. Large systematic errors of -90 to 300% were found when it was assumed that sap flow was uniform over the sapwood depth. Therefore, we recommend that the radial sap flow pattern should be determined first using sensors with multiple measuring points along a stem radius followed by single-point measurements with sensors placed at a predetermined depth. Other significant errors occurred in the scaling procedure even when the sap flow radial pattern was known. These included errors associated with uncertainties in the positioning of sensors beneath the cambium (up to 15% per 1 mm error in estimated xylem depth), and differences in environmental conditions when the radial profile applied for integration was determined over the short term (up to 47% error). High temporal variation in the point-to-area correction factor along the xylem radius used for flow integration is also problematic. Compared with midday measurements, measurements of radial variation of sap flow in the morning and evening of sunny days minimized the influence of temporal variations on the point-to-area correction factor, which was especially pronounced in trees with a highly asymmetric sap flow radial pattern because of differences in functioning of the sapwood xylem layers. Positioning a single-point sensor at a depth with maximum sap flow is advantageous because of the high sensitivity of maximum sap flow to water stress conditions and changes in micro-climate, and because of the lower random errors associated with the positioning of a single-point sensor along the xylem radius.     

Variation in the radial patterns of sap flux density in pubescent oak (Quercus pubescens) and its implications for tree and stand transpiration measurements

Radial variation in sap flux density across the sapwood was assessed by the heat field deformation method in several trees of Quercus pubescens Wild., a ring-porous species. Sapwood depths were delimited by identifying the point of zero flow in radial patterns of sap flow, yielding tree sapwood areas that were 1.5-2 times larger than assumed based on visual examinations of wood cores. The patterns of sap flow varied both among trees and diurnally. Rates of sap flow were higher close to the cambium, although there was a significant contribution from the inner sapwood, which was greater (up to 60% of total flow) during the early morning and late in the day. Accordingly, the normalized difference between outer and inner sapwood flow was stable during the middle of the day, but showed a general decline in the afternoon. The distribution of sap flux density across the sapwood allowed us to derive correction coefficients for single-point heat dissipation sap flow measurements. We used daytime-averaged coefficients that depended on the particular shape of the radial profile and ranged between 0.45 and 1.28. Stand transpiration calculated using the new method of estimating sapwood areas and the radial correction coefficients was similar to (Year 2003), or about 25% higher than (Year 2004), previous uncorrected values, and was 20-30% of reference evapotranspiration. We demonstrated how inaccuracies in determining sapwood depths and mean sap flux density across the sapwood of ring-porous species could affect tree and stand transpiration estimates.     

Integration of sapflow velocity in elliptical stems

The estimation of water flux through the stem of a plant by point estimates based on the heat pulse technique requires integration of the velocity profile. The polynomial integration method and the weighted average technique both assume radial symmetry about the bole. A method is introduced that solves the equations for the ellipse in the weighted average technique, and includes an optimizing algorithm for the placement of sensors. The sensitivity of sapflow estimates through stems to increasing eccentricity is examined by means of analytical solutions and simulation. The error in sap flux measurement resulting from the radial approximation of the sapwood conducting area is expressed analytically and shown to be small over the range of eccentricities (e < 0.8) expected in nature. Simulations where radial symmetry is assumed for elliptical stems show substantial error in mean velocity (and thus flux) with modest eccentricities (e > 0.4).     

Radial profiles of sap flow with increasing tree size in maritime pine

We investigated the radial variation of sap flow within sapwood below the live crown in relation to tree size in 10-, 32-, 54- and 91-year-old maritime pine stands (Pinus pinaster Ait.). Radial variations were determined with two thermal dissipation sensors; one measured sap flux in the outer 20 mm of the xylem (Jref), whereas the other was moved radially across the sapwood in 20-mm increments to measure sap flux at multiple depths (Jref). For all tree sizes, sap flow ratios (Ri = JiJref (-1)) declined with increasing sapwood depth, but the decrease was steeper in trees with large diameters. Correction factors (C) were calculated to extrapolate Jref for an estimate of whole-tree sap flux. A negative linear relationship was established between stem diameter and C, the latter ranging from 0.6 to 1.0. We found that neglecting these radial corrections in 10-, 32-, 54- and 91-year-old trees would lead to overestimation of stand transpiration by 4, 14, 26 and 47%, respectively. Therefore, it is necessary to account for the differential radial profiles of sap flow in relation to tree size when comparing tree transpiration and hydraulic properties among trees differing in size.     

Medium-term sap flux monitoring in a Scots pine stand: analysis of the operability of the heat dissipation method for hydrological purposes

The operability of Granier-type heat dissipation sap flow meters for the medium-term monitoring of Scots pine transpiration was tested. Three sensors that had been working for 3 years were duplicated by inserting new sensors in the same trees. The new sensors operated simultaneously with the old sensors for 18 months. Analysis of the temporal patterns of thermal dissipation of the sensors showed a slight, but significant decrease in all sensors, indicating the conservation of sapwood thermal properties. Although there was a high correlation between sap flux densities registered by the old and new sensors, significant differences in sap flux densities between the duplicated sensors were detected. Such differences could not be attributed to tissue injury around the sensors or to loss of sensitivity of the old sensors, because two of the old sensors registered higher flux rates than the new sensors. No instrumental error to explain the sap flux differences was found. Because the new sensors were installed at some angular distance from the old sap flow meters to avoid thermal interferences, it was inferred that the observed sap flux differences between duplicate sensors were the result of an azimuthal sap flow pattern.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

A model of heat transfer in sapwood and implications for sap flux density measurements using thermal dissipation probes

A variety of thermal approaches are used to estimate sap flux density in stems of woody plants. Models have proved valuable tools for interpreting the behavior of heat pulse, heat balance and heat field deformation techniques, but have seldom been used to describe heat transfer dynamics for the heat dissipation method. Therefore, to better understand the behavior of heat dissipation probes, a model was developed that takes into account the thermal properties of wood, the physical dimensions and thermal characteristics of the probes, and the conductive and convective heat transfer that occurs due to water flow in the sapwood. Probes were simulated as aluminum tubes 20 mm in length and 2 mm in diameter, whereas sapwood, heartwood and bark each had a density and water fraction that determined their thermal properties. Base simulations assumed a constant sap flux density with sapwood depth and no wounding or physical disruption of xylem beyond the 2 mm diameter hole drilled for probe installation. Simulations across a range of sap flux densities showed that the dimensionless quantity k [defined as (ΔT(m) -ΔT)/ΔT, where ΔT(m) is the temperature differential (ΔT) between the heated and unheated probe under zero-flow conditions] was dependent on the thermal conductivity of the sapwood. The relationship between sap flux density and k was also sensitive to radial gradients in sap flux density and to xylem disruption near the probe. Monte Carlo analysis in which 1000 simulations were conducted while simultaneously varying thermal conductivity and wound diameter revealed that sap flux density and k showed considerable departure from the original calibration equation used with this technique. The departure was greatest for variation in sap flux density typical of ring-porous species. Depending on the specific combination of thermal conductivity and wound diameter, use of the original calibration equation resulted in an 81% under- to 48% overestimation of sap flux density at modest flux rates. Future studies should verify these simulations and assess their utility in estimating sap flux density for this widely used technique.     

Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.     

Estimating water use by sugar maple trees: considerations when using heat-pulse methods in trees with deep functional sapwood

Accurate estimates of sapwood properties (including radial depth of functional xylem and wood water content) are critical when using the heat pulse velocity (HPV) technique to estimate tree water use. Errors in estimating the volumetric water content (V(h)) of the sapwood, especially in tree species with a large proportion of sapwood, can cause significant errors in the calculations ofsap velocity and sap flow through tree boles. Scaling to the whole-stand level greatly inflates these errors. We determined the effects of season, tree size and radial wood depth on V(h) of wood cores removed from Acer saccharum Marsh. trees throughout 3 years in upstate New York. We also determined the effects of variation in V(h) on sap velocity and sap flow calculations based on HPV data collected from sap flow gauges inserted at four depths. In addition, we compared two modifications of Hatton's weighted average technique, the zero-step and zero-average methods, for determining sap velocity and sap flow at depths beyond those penetrated by the sap flow gauges. Parameter V(h) varied significantly with time of year (DOY), tree size (S), and radial wood depth (RD), and there were significant DOY x S and DOY x RD interactions. Use of a mean whole-tree V(h) value resulted in differences ranging from -6 to +47% for both sap velocity and sap flow for individual sapwood annuli compared with use of the V(h) value determined at the specific depth where a probe was placed. Whole-tree sap flow was 7% higher when calculated on the basis of the individual V(h) value compared with the mean whole-tree V(h) value. Calculated total sap flow for a tree with a DBH of 48.8 cm was 13 and 19% less using the zero-step and the zero-average velocity techniques, respectively, than the value obtained with Hatton's weighted average technique. Smaller differences among the three methods were observed for a tree with a DBH of 24.4 cm. We conclude that, for Acer saccharum: (1) mean V(h) changes significantly during the year and can range from nearly 50% during winter and early spring, to 20% during the growing season;(2) large trees have a significantly greater V(h) than small trees; (3) overall, V(h) decreases and then increases significantly with radial wood depth, suggesting that radial water movement and storage are highly dynamic; and (4) V(h) estimates can vary greatly and influence subsequent water use calculations depending on whether an average or an individual V(h) value for a wood core is used. For large diameter trees in which sapwood comprises a large fraction of total stem cross-sectional area (where sap flow gauges cannot be inserted across the entire cross-sectional area), the zero-average modification of Hatton's weighted average method reduces the potential for large errors in whole-tree and landscape water balance estimates based on the HPV method.     

Forest transpiration—targeted through xylem sap flux assessment versus hydrological modeling

The assessment of forest transpiration rates is crucial for determining plant-available soil water consumption and drought risk of trees. Xylem sap flux measurements have been used increasingly to quantify stand transpiration in forest ecosystems. Here, we compare this empirical approach with hydrological modeling on the basis of a stand transpiration dataset of adult beech (Fagus sylvatica), which was acquired across Bavaria, Germany, at eight forest sites. Xylem sap flux sensors were installed in five dominant trees each. Two tree to stand upscaling approaches, related to site-specific (1) sapwood area or (2) to leaf area index, were compared. The outcome was examined each in relation to process-based stand hydrological modeling, using LWF-BROOK90. Distinct relationships between tree diameter at breast height (1.30 m) and sapwood area-weighted sap flux along the radial profile became apparent across the study sites, confirming a generic allometric basis for stand-level upscaling of transpiration. The two upscaling approaches did not differ in outcome, representatively covering stand structure for comparison with modeling. Differential analysis yielded high agreement between the empirical and modeling approaches throughout most of the study period, although LWF-BROOK90 tended to overestimate sap flux measurements under low soil moisture. The two empirical approaches proved reliable for even-aged beech stands, as performance under high stand-structural heterogeneity awaits clarification. Findings advance stand-level hydrological modeling regarding coverage of stomatal behavior during temporary limitation in water availability.     

Vertical foliage distribution determines the radial pattern of sap flux density in Picea abies

Understanding the causes determining the radial pattern of sap flux density is important both for improving knowledge of sapwood functioning and for up-scaling sap flow measurements to canopy transpiration and ecosystem water use. To investigate the anatomical connection between whorls and annual sapwood rings, pruning-induced variation in the radial pattern of sap flux density was monitored with Granier probes in a 35-year-old Picea abies (L.) Karst tree that was pruned from the crown bottom up. Modifications in the radial pattern of sap flux density were quantified by a shape index (SI), which varies with the relative contribution of the outer and inner sapwood to tree transpiration. The SI progressively diminished during bottom up pruning, indicating a significant reduction in sap flow contribution of the inner sapwood. Results suggest that the radial pattern of sap flux density depends mainly on the vertical distribution of foliage in the crown, with lower shaded branches hydraulically connected with inner sapwood and upper branches connected with the outer rings.     

桉树液流径向变化及其对整树蒸腾估算的影响

目的 分析液流密度径向差异及其对整树蒸腾量估算的影响,以期为中大径级桉树人工林蒸腾的准确估算和尺度扩展提供可靠依据。 方法 采用Granier热扩散探针法(TDP)测定了10年生中等径级尾巨桉(Eucalyptus urophylla × Eucalyptus grandis)3个边材径向深度0～2、2～4、4～6 cm的液流密度。 结果 尾巨桉液流密度径向分布格局为递减型,日变化格局均为单峰型。用单点径向深度0～2、2～4、4～6 cm液流密度代表整个边材液流密度估算的整树日蒸腾量Tr2、Tr4和Tr6较基于多点估算值Tr246分别高估92.9%、低估28.2%和低估74.0%。由于0～2 cm 边材深度液流密度对环境变化较为敏感,并且 Tr2与Tr246之间的拟合精度较高(R2=0.964),因此,基于Tr2构建的整树日蒸腾量估算校正函数Tr246=0.522Tr2有效的提高了整树蒸腾估算的便捷性和准确度。 结论 中等径级尾巨桉液流密度具有明显的径向变化特征,不考虑这种径向变化将会导致整树蒸腾估算的较大误差。基于边材最外侧（0～2 cm）液流密度估算的整树蒸腾经过校正后可以方便准确地估算中等径级桉树人工林蒸腾耗水。     

Potential errors in measurement of nonuniform sap flow using heat dissipation probes

The empirical calibration of Granier-type heat dissipation sap flow probes that relate temperature difference (DeltaT) to sap velocity (v) was reevaluated in stems of three tropical tree species. The original calibration was confirmed when the entire heated probe was in contact with conducting xylem, but mean v was underestimated when part of the probe was in contact with nonconducting xylem or bark. Analysis of the effects of nonuniform sap velocity profiles on heat dissipation estimates showed that errors increased as v and the proportion of the probe in nonconducting wood increased. If half of a 20-mm probe is in sapwood with a v of 0.15 mm s(-1) and the other half is in nonconducting wood, then mean v for the whole probe can be underestimated by as much as 50%. A correction was developed that can be used if the proportion of the probe in nonconducting wood is known. Even with the entire heated probe in contact with conducting xylem, v would be underestimated when radial velocity gradients are present. In this case, the error would be smaller except when velocity gradients are very steep, as can occur in species with ring-porous wood anatomy. Errors occur because the relationship between DeltaT and v is nonlinear. Mean DeltaT along the probe is therefore not a measure of mean v, and users of heat dissipation probes should not assume that v is integrated along the length of the probe. The same type of error can occur when DeltaT is averaged through time while v is changing, but the error is small unless there are sudden, step changes between zero and high sap velocity. It is recommended that relatively short probes (20 mm or less) be used and that probes longer than the depth of conducting sapwood be avoided. Multiple probes inserted to a range of depths should be used in situations where steep gradients in v are expected. If these conditions are met, heat dissipation probes remain useful and widely applicable for measuring sap flow in woody stems.     

Regulation of water flux through tropical forest canopy trees: do universal rules apply?

Tropical moist forests are notable for their richness in tree species. The presence of such a diverse tree flora presents potential problems for scaling up estimates of water use from individual trees to entire stands and for drawing generalizations about physiological regulation of water use in tropical trees. We measured sapwood area or sap flow, or both, in 27 co-occurring canopy species in a Panamanian forest to determine the extent to which relationships between tree size, sapwood area and sap flow were species-specific, or whether they were constrained by universal functional relationships between tree size, conducting xylem area, and water use. For the 24 species in which active xylem area was estimated over a range of size classes, diameter at breast height (DBH) accounted for 98% of the variation in sapwood area and 67% of the variation in sapwood depth when data for all species were combined. The DBH alone also accounted for > or = 90% of the variation in both maximum and total daily sap flux density in the outermost 2 cm of sapwood for all species taken together. Maximum sap flux density measured near the base of the tree occurred at about 1,400 h in the largest trees and 1,130 h in the smallest trees studied, and DBH accounted for 93% of the variation in the time of day at which maximum sap flow occurred. The shared relationship between tree size and time of maximum sap flow at the base of the tree suggests that a common relationship between diurnal stem water storage capacity and tree size existed. These results are consistent with a recent hypothesis that allometric scaling of plant vascular systems, and therefore water use, is universal.     

Enhanced transpiration in response to wind effects at the edge of a blue gum (Eucalyptus globulus) plantation

In Australia, tree planting has been widely promoted to alleviate dryland salinity and one proposed planting configuration is that of strategically placed interception belts. We conducted an experiment to determine the effect of tree position in a belt on transpiration rate. We also assessed how much the effect of tree position can be explained by advection and environmental conditions. Daily transpiration rates were determined by the heat pulse velocity technique for four edge and 12 inner trees in a 7-year-old Tasmanian blue gum (Eucalyptus globulus) plantation in South Australia. Various climatic variables were logged automatically at one edge of the plantation. The relationship between daily sap flow and sapwood area was strongly linear for the edge trees (r2 = 0.97), but only moderately correlated for the inner trees (r2 = 0.46), suggesting an edge effect. For all trees, sap flow normalized to sapwood area (Qs) increased with potential evaporation (PE) initially and then became independent as PE increased further. There was a fairly close correlation between transpiration of the edge and inner trees, implying that water availability was partially responsible for the difference between inner and edge trees. However, the ratio of edge tree to inner tree transpiration differed from unity, indicating differences in canopy conductance, which were estimated by an inverse form of the Penman-Monteith equation. When canopy conductances were less than a critical value, there was a strong linear relationship between Qs of the edge and inner trees. When canopy conductances of the edge trees were greater than the critical value, the slope of the linear relationship was steeper, indicating greater transpiration of the edge trees compared with the inner trees. This was interpreted as evidence for enhancement of transpiration of the edge trees by advection of wind energy.     

Spatial variations in xylem sap flux density in the trunk of orchard-grown,mature mango trees under changing soil water conditions

Circumferential and radial variations in xylem sap flux density in trunks of 13-year-old mango (Mangifera indica L.) trees were investigated with Granier sap flow sensor probes under limiting and non-limiting soil water conditions. Under non-limiting soil water conditions, circumferential variation was substantial, but there was no apparent relationship between sap flux density and aspect (i.e., the radial position of the sensor probes on the trunk relative to the compass). Hourly sap flux densities over 24 hours at different aspects were highly pair-wise correlated. The relationships between different aspects were constant during well-watered periods but highly variable under changing soil water conditions. Sap flux density showed marked radial variation within the trunk and a substantial flux was observed at the center of the trunk. For each selected aspect on each tree, changes in sap flux densities over time at different depths were closely correlated, so flux at a particular depth could be extrapolated as a multiple of flux from 0 to 2 cm beneath the cambium. However, depth profiles of sap flux density differed between trees and even between aspects within a tree, and also varied in an unpredictable manner as soil water conditions changed. Nevertheless, over a period of non-limiting soil water conditions, depth profiles remained relatively constant. Based on the depth profiles obtained during these periods, a method is described for calculating total sap flow in a mango tree from sap flux density at 0-2 cm beneath the cambium. Total daily sap flows obtained were consistent with water use estimated from soil water balance.