Assessing variation in the radial profile of sap flux density in Pinus species and its effect on daily water use

We monitored sap flux density (v) diurnally in nine mature southeastern pine (Pinus spp.) trees with a thermal dissipation probe that spanned the sapwood radius. We found the expected pattern of high v near the cambium and decreasing v with depth toward the center of the tree; however, the pattern was not constant within a day or between trees. Radial profiles of trees were steeper earlier in the day and became less steep later in the day. As a result, time-dependent changes in the shape of the radial profile of v were sometimes correlated with daily changes in evaporative demand. As the radial profile became less steep, the inner xylem contributed relatively more to total tree sap flow than it did earlier in the day. We present a 3-parameter Gaussian function that can be used to describe the radial distribution of v in trees. Parameters in the function represent depth in the xylem from the cambium, maximum v, depth in the xylem where maximum v occurs, and the rate of radial change in v with radial depth (beta). Values of beta varied significantly between trees and with time, and were sometimes correlated with air vapor pressure deficit (D). We hypothesize that this occurred during periods of high transpiration when the water potential gradient became great enough to move water in the inner sapwood despite its probable high hydraulic resistance. We examined discrepancies among estimates of daily water use based on single-point, two-point and multi-point (i.e., every 20 mm in the sapwood) measurements. When radial distribution of v was not considered, a single-point measurement resulted in errors as large as 154% in the estimate of daily water use relative to the estimate obtained from a multi-point measurement. Measuring v at two close sample points (10 and 30 mm) did not improve the estimate; however, estimates derived from v measured at two distant sample points (10 and 70 mm) significantly improved the estimate of daily water use, although errors were as great as 32% in individual trees. The variability in v with depth in the xylem, over time, and between trees indicates that measurements of the radial distribution of v are necessary to accurately estimate water flow in trees with large sapwood areas.     

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.     

Radial variation in sap flow in five laurel forest tree species in Tenerife,Canary Islands

Variations in radial patterns of xylem water content and sap flow rate were measured in five laurel forest tree species (Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng., Myrica faya Ait., Erica arborea L. and Ilex perado Ait. ssp. platyphylla (Webb & Berth.) Tutin) growing in an experimental plot at Agua García, Tenerife, Canary Islands. Measurements were performed around midday during warm and sunny days by the heat field deformation method. In all species, water content was almost constant (around 35% by volume) over the whole xylem cross-sectional area. There were no differences in wood color over the whole cross-sectional area of the stem in most species with the exception of E. arborea, whose wood became darker in the inner layers. Radial patterns of sap flow were highly variable and did not show clear relationships with tree diameter or species. Sap flow occurred over the whole xylem cross-sectional area in some species, whereas it was limited to the outer xylem layers in others. Sap flow rate was either similar along the xylem radius or exhibited a peak in the outer part of the xylem area. Low sap flow rates with little variation in radial pattern were typical for shaded suppressed trees, whereas dominant trees exhibited high sap flow rates with a peak in the radial pattern. Stem damage resulted in a significant decrease in sap flow rate in the outer xylem layers. The outer xylem is more important for whole tree water supply than the inner xylem because of its larger size. We conclude that measurement of radial flow pattern provides a reliable method of integrating sap flow from individual measuring points to the whole tree.     

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.     

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.     

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.     

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.     

Radial distribution of sap flux density in trunks of a mature beech stand

In a mature beech stand located in north-eastern Germany, xylem sap flux measurements were continuously performed during the 2002–2004 growing seasons. Ten representative trunks were studied using heated thermal dissipation probes. The measurements aimed at identifying principles governing radial profiles of xylem flux in beech trunks. The measurements were taken up to a trunk depth of 132 mm. The sap flow density in the pericambial xylem was found to vary among trees of different diameters, but was not considerably smaller in suppressed trees. A model for the radial distribution of sap flux density was formulated relating trunk radius and sap flow density. The model takes into account different trunk diameter. About 90% of the sap flux was found to occur in the outer two fifths of the trunk. Using this model, an adequate estimate of transpiration can be achieved at tree and stand level, even when the sap flux measurements are restricted to the outer trunk sectors.     

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.     

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.     

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.     

Sap flow characteristics of neotropical mangroves in flooded and drained soils

Effects of flooding on water transport in mangroves have previously been investigated in a few studies, most of which were conducted on seedlings in controlled settings. In this study, we used heat-dissipation sap probes to determine if sap flow (J(s)) attenuates with radial depth into the xylem of mature trees of three south Florida mangrove species growing in Rookery Bay. This was accomplished by inserting sap probes at multiple depths and monitoring diurnal flow. For most species and diameter size class combinations tested, J(s) decreased dramatically beyond a radial depth of 2 or 4 cm, with little sap flow beyond a depth of 6 cm. Mean J(s) was reduced on average by 20% in Avicennia germinans (L.) Stearn, Laguncularia racemosa (L.) Gaertn. f. and Rhizophora mangle L. trees when soils were flooded. Species differences were highly significant, with L. racemosa having the greatest midday J(s) of about 26 g H(2)O m(-2) s(-1) at a radial depth of 2 cm compared with a mean for the other two species of about 15 g H(2)O m(-2) s(-1). Sap flow at a depth of 2 cm in mangroves was commensurate with rates reported for other forested wetland tree species. We conclude that: (1) early spring flooding of basin mangrove forests causes reductions in sap flow in mature mangrove trees; (2) the sharp attenuations in J(s) along the radial profile have implications for understanding whole-tree water use strategies by mangrove forests; and (3) regardless of flood state, individual mangrove tree water use follows leaf-level mechanisms in being conservative.     

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 water transport pathways in a maple tree: responses of sap flow to branch severing

? It has been known for a long time that sectored and integrated patterns of vascular systems exist in different species and even within the same tree, depending on its age and history. However, very few publications consider the topology of the vascular pathways between roots and branches.

? Some results on this important aspect of the vascular system are presented in this paper. They have been obtained with adult maple trees by directly studying the water movement in the stem and root xylem with the heat field deformation (HFD) method for sap flow measurements.

? Multi-point HFD sensors were installed at different heights of a Norway maple tree (Acer platanoides L.) along its stem axis. Single-point HFD sensors were installed in three small lateral roots of another sample maple. Experimental treatments (branch severing) triggered changes in sap movement in the stem and root sapwood.

? The sample trees belong to the group with an integrated transport system (“integrated pipes”), sharing stem space on both sides of the tree to supply two large parts of the crown with water from each root sector. Nevertheless, conducting pathways had their autonomy for axial transport and the pipe model theory describes the vascular system of the studied trees well. Thus, the integration of axial transport in the stem xylem should presumably occur through the cross-grained network of axial vessels.

     

Calibration of sap flow estimated by the compensation heat pulse method in olive, plum and orange trees: relationships with xylem anatomy

The compensation heat pulse method is widely used to estimate sap flow in conducting organs of woody plants. Being an invasive technique, calibration is crucial to derive correction factors for accurately estimating the sap flow value from the measured heat pulse velocity. We compared the results of excision and perfusion calibration experiments made with mature olive (Olea europaea L. 'Manzanilla de Sevilla'), plum (Prunus domestica L. 'Songal') and orange (Citrus sinensis (L.) Osbeck. 'Cadenero') trees. The calibration experiments were designed according to current knowledge on the application of the technique and the analysis of measured heat pulse velocities. Data on xylem characteristics were obtained from the experimental trees and related to the results of the calibration experiments. The most accurate sap flow values were obtained by assuming a wound width of 2.0 mm for olive and 2.4 mm for plum and orange. Although the three possible methods of integrating the sap velocity profiles produced similar results for all three species, the best results were obtained by calculating sap flow as the weighted sum of the product of sap velocity and the associated sapwood area across the four sensors of the heat-pulse-velocity probes. Anatomical observations showed that the xylem of the studied species can be considered thermally homogeneous. Vessel lumen diameter in orange trees was about twice that in the olive and plum, but vessel density was less than half. Total vessel lumen area per transverse section of xylem tissue was greater in plum than in the other species. These and other anatomical and hydraulic differences may account for the different calibration results obtained for each species.     

Measured and predicted changes in tree and stand water use following high-intensity thinning of an 8-year-old Eucalyptus nitens plantation

We investigated changes in the pattern of water use of an 8-year-old Eucalyptus nitens (Deane and Maiden) Maiden plantation soon after thinning. Sap flow sensors using heat pulse technology were deployed across three stands thinned to a final density of 100, 250 or 600 trees ha-1 plus an unthinned control (1250 trees ha-1). Changes in the relationship between tree size and daily water use were measured for 4 to 7 months after thinning. Thinning had no effect on sapwood water content. The increase in tree water use as a result of thinning was driven largely by significant changes in the radial pattern of sap velocity through the sapwood. The use of a canopy fraction factor in the Penman-Monteith equation to account for discontinuous canopies showed promise as a simple and effective method of scaling the model to predict transpiration from thinned plantations.     

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.     

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

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

Variation of sapflow velocity in Eucalyptus globulus with position in sapwood and use of a correction coefficient

Sapflow sensors were used to investigate variation in sapflow velocity at different positions in the sapwood of three-year-old Eucalyptus globulus ssp. globulus Labill. trees. Sapflow velocity was measured at 5-mm intervals across the sapwood by moving two probe sets simultaneously on two opposite radii. Another probe set placed in a fixed position at right angles to the first two sets acted as a control. A sapflow velocity ratio was defined as the velocity given by each moving sensor divided by that given by the static sensor. Correlation between observations of sapflow velocity at different positions exceeded 95%, and the ratio of velocity between any pair of sensors was constant. We observed radial variation in sapflow velocity across the sapwood with the lowest velocities at the center of the tree. Variation due to sensor position was high implying the need for large numbers of sensors for accurate estimates of sap flux. To overcome this need, we used a correction coefficient, namely a simple weighted average of the sapflow ratios with depth in the sapwood, for each fixed sensor. We recommend the use of three probe sets to estimate the correction coefficient. Subsequently, two probe sets can be placed at two fixed positions for routine measurements of sap flux.     

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.