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.     

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.     

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.     

Influences of environmental factors on the radial profile of sap flux density in Fagus crenata growing at different elevations in the Naeba Mountains, Japan

Sap flux density was measured continuously during the 1999 and 2000 growing seasons by the heat dissipation method in natural Fagus crenata Blume (Japanese beech) forests growing between 550 and 1600 m on the northern slope of the Kagura Peak of the Naeba Mountains, Japan. Sap flux density decreased radially toward the inner xylem and the decrease was best expressed in relation to the number of annual rings from the cambium, or in relation to the relative depth between the cambium and the trunk center, rather than as a function of absolute depth. The relative influences of radiation, vapor pressure deficit and soil water on sap flux density during the growing season were similar for the outer and inner xylem, and at all sites. Measurements of soil water content and water potential at a depth of 0.25 m demonstrated that sap flux density responded similarly and sensitively to water potential changes in this soil layer, despite large differences in rooting depth at different elevations, localizing one important control point in the functioning of this forest ecosystem. Identification of the relative influences of radiation, vapor pressure deficit and drying of the upper soil layer on sap flux density provides a framework for in-depth analysis of the control of transpiration in Japanese beech forests. In addition, the finding that the same general controls are operating on sap flux density despite climate gradients and large differences in overall forest stand structure will enhance understanding of water use by forests along elevation gradients.     

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.     

Seasonal changes in the axial distribution of peroxidase activity in the xylem sap of beech (Fagus sylvatica L.) trees

Xylem sap was collected from trunk segments of adult beech (Fagus sylvatica L.) trees by water displacement. Peroxidase activity was analyzed in xylem saps collected in different phases of the yearly growth cycle and from different heights up the trunks (up to 14 m). The xylem saps contained two major peroxidase isozymes with acidic isoelectric points of 4.1 and 4.6, respectively. Mean peroxidase activity was low during the emergence of the new leaves and high in summer and in winter. In the cold season, peroxidase activity decreased from the stem base to the top, whereas significant gradients were not observed during the vegetative period.     

Xylem sap composition of beech (Fagus sylvatica L.) trees: seasonal changes in the axial distribution of sulfur compounds

During different phases of the annual growth cycle, xylem sap was collected from trunk segments of adult beech (Fagus sylvatica L.) trees by the water displacement technique. Irrespective of the height of the trunk, both sulfate and reduced sulfur compounds were detected in the xylem sap throughout the year. Sulfate was the predominant sulfur compound in all samples analyzed. Its concentration in the xylem sap varied between 10 and 350 micro mol l(-1), with highest concentrations in April, shortly before bud break. In contrast to other tree species, cysteine and not glutathione was the predominant thiol transported in the xylem sap of beech trees. The cysteine concentration ranged between 0.1 and 1 micro mol l(-1). As observed for sulfate, maximum cysteine concentrations were found in April. Apparently, both sulfate and cysteine transport contribute to the sulfur supply of the developing leaves. Seasonal changes in the axial distribution of cysteine and sulfate differed, indicating differences in the source-sink relations of these sulfur compounds. High, but uniform, xylem sap sulfate concentrations in April may originate from balanced sulfate uptake by the roots, whereas high cysteine concentrations in April, increasing with increasing height of the trunk, may originate in part from protein breakdown in the trunk. Reversal of the axial distribution of xylem sap cysteine in late summer-early fall to higher concentrations in the lower part of the trunk than in the upper part of the trunk suggests that the upper part of the trunk becomes a sink for cysteine as a result of the synthesis of storage proteins at this time of the year.     

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.     

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.     

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.     

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.     

Axial and radial water flow in the trunks of oak trees: a quantitative and qualitative analysis

Axial water flow in the trunks of mature oak trees (Quercus petraea (Matt.) Liebl. and Q. robur L.) was studied by four independent techniques: water absorption from a cut trunk, sap flowmeters, heat pulse velocity (HPV) and thermoimaging. Estimation of the total water flow with sap flowmeters, HPV and water absorption yielded comparable results. We concluded from dye colorations, thermograms and axial profiles of sap flow and heat pulse velocity that, in intact trunks, most of the flow occurred in the current-year ring, where early-wood vessels in the outermost ring were still functional. Nevertheless, there was significant flow in the older rings of the xylem. Total water flow through the trunk was only slightly reduced when air embolisms were artificially induced in early-wood vessels, probably because there was little change in hydraulic conductance in the root-leaf sap pathway. Embolization of the current-year vessels reactivated transport in the older rings.     

Sap-flux-scaled transpiration responses to light,vapor pressure deficit,and leaf area reduction in a flooded Taxodium distichum forest

We used 20-mm-long, Granier-type sensors to quantify the effects of tree size, azimuth and radial position in the xylem on the spatial variability in xylem sap flux in 64-year-old trees of Taxodium distichum L. Rich. growing in a flooded forest. This information was used to scale flux to the stand level to investigate variations in half-hourly and daily (24-hour) sums of sap flow, transpiration per unit of leaf area, and stand transpiration in relation to vapor pressure deficit (D) and photosynthetically active radiation (Q(o)). Measurements of xylem sap flux density (J(s)) indicated that: (1) J(s) in small diameter trees was 0.70 of that in medium and large diameter trees, but the relationship between stem diameter as a continuous variable and J(s) was not significant; (2) J(s) at 20-40 mm depth in the xylem was 0.40 of that at 0-20 mm depth; and (3) J(s) on the north side of trees was 0.64 of that in directions 120 degrees from the north. Daily transpiration was linearly related to daily daytime mean D, and reached a modest value of 1.3 mm day(-1), reflecting the low leaf area index (LAI = 2.2) of the stand. Because there was no soil water limitation, half-hourly water uptake was nearly linearly related to D at D < 0.6 kPa during both night and day, increasing to saturation during daytime at higher values of D. The positive effect of Q(o) on J(s) was significant, but relatively minor. Thus, a second-order polynomial with D explained 94% of the variation in J(s) and transpiration. An approximately 40% reduction in LAI by a hurricane resulted in decreases of about 18% in J(s) and stand transpiration, indicating partial stomatal compensation.     

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.     

Transpiration of Scots pine in Flanders growing on soil with irregular substratum

An investigation was carried out to compare the water balance of Scots pine in Flanders growing on soils with contrasted water availability. Based on sap flow measurements transpiration of Scots pine was determined for two small plots on cover sands resting on a clayey substratum of varying depths (shallow and deep). Soil water content (SWC) was relatively low (0.12–0.21 m³ m⁻³) in the upper topsoil (0–0.75 m) in both plots. However, it was always higher in the shallow plot (by 3–27%) than in the deep plot. The difference between SWC in both plots was more pronounced in the deeper soil layers (0.75–1.5 m). Sap flow was measured in seven sample pine trees on each plot from May to October 2000 using the heat field deformation (HFD) method. Transpiration of the individual trees in the deep plot was 22% lower than in trees in the shallow plot. The difference decreased to 15% after scaling up to the stand level due to a higher density of trees growing in the deep plot. It was hypothesized that higher water uptake in the shallow plot was possibly caused by structural differences between the root systems of trees growing in plots with variable soil texture. The sapwood in shallow-plot trees was 1 cm less deep than in trees growing in the deep plot (as measured by biometric and sap flow pattern methods). Sap flow radial patterns suggested a higher involvement of sinker roots for water uptake in the deep clayey substratum plot. This was in agreement with higher activity of the inner xylem in trees on the deep plot under higher evaporative demands. However, the fraction of the inner xylem to the whole-tree water supply was nearly three-fold lower than the outer xylem, which appeared to provide water presumably from the superficial roots. The fraction of these roots, estimated according to sap flow radial patterns, was around 10% higher in trees on the shallow plot. This caused 30% higher sap flow in the stem outer xylem there. Transpiration of the pine stands was limited under high evaporative demands in both plots by the low availability of soil water. The limitation was greater in the deep plot and persisted throughout the whole growing season.     

Growth strain in coconut palm trees

Until recently, growth stress studies have been made only on coniferous and dicotyledonous trees. Growth stress of trees is thought to be initiated in newly formed secondary xylem cells. This stress can accumulate for years and is distributed inside the trunk. Major characteristics of the trunk of monocotyledonous trees include numerous vascular bundles scattered inside the ground tissue and the lack of secondary growth for enlarging the diameter of the trunk. We used the strain gauge method to measure the released growth strain of the monocotyledonous woody palm, coconut (Cocos nucifera L.), and to investigate the surface growth strain of the trunk and central cylinder at different trunk heights. The internal strains of both vertical and leaning trunks were measured and compared with those of coniferous and dicotyledonous trees. We found that tensile stress existed longitudinally on the surface of vertically growing trunks, whereas compression stress was found at the bending position of leaning trunks. Compression stress was found in the outer part of the central cylinder, whereas tensile stress is generally found in the outer part of the trunk in coniferous and dicotyledonous trees. The distribution of strain in the palm trunk is similar to that of compression wood of the leaning trunk of a conifer. Specific gravity was greater in the outer part of the trunk than in the inner part of the trunk. This difference may be related to the distribution of growth stress.     

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.     

Radial distribution of carbohydrate reserves in the trunk of declining European beech trees (Fagus sylvatica L.)

Context

European beech (Fagus sylvatica L.) is considered threatened by anticipated climate change, but the physiological causes of potential beech decline or mortality remains poorly understood.

Aims

The purpose of the present study was to fuel debate about the assumption that carbohydrate depletion is involved in the decline of mature European beech.

Methods

The health status of beech trees from a severely declining stand was visually assessed by examining their crown condition. Content and radial distribution of non-structural carbohydrates (starch and soluble carbohydrate) were analyzed in the trunks and compared to those reported earlier in trunks of healthy beech trees.

Results and discussion

The distribution of carbohydrate in the beech trunks recorded here seemed affected by decline. We found a stronger radial decrease of starch content than those reported earlier for healthy beech trees. Carbohydrate reserves appear partially maintained in the outermost rings while starch depletion occurred in older wood rings in declining trees that may be able to mobilize carbohydrate reserves from older wood rings in response to successive climatic constraints.