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

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

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.     

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

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

Stem sapwood permeability in relation to crown dominance and site quality in self-thinning fire-origin lodgepole pine stands

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.     

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

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

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.     

Dependence of shear strength on wood properties in cultivated Larix sibirica

Abstract

Mechanical properties are very important in structural uses of wood. In the case of larch wood, shear strength (SS) may be a limiting property, because the wood is brittle and it splinters easily. In this study, SS of Siberian larch (Larix sibirica) wood was studied in relation to its axial and radial location and to some other wood properties in the part of the trunk that produces logs of good carpentry quality. SS increased by 21–23% from the pith to the outer heartwood at each studied height (butt, 4.5 and 9 m), and it was at its most approximately 7% higher at the butt than at the other studied heights. The main reason for the radial increase in SS was the increase in density caused by wood maturation, but other factors also affected SS, depending on the grouping mode (combined material, radial/axial location). An exception was sapwood, for which no factors affecting SS could be identified. Thus, sapwood may behave differently from heartwood regarding use, even unexpectedly. The results suggest that mature heartwood, but not juvenile wood and sapwood, of Larix sibirica should primarily be used in products that require high SS.     

Changes in sapwood permeability and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans

Increases in plant size and structural complexity with increasing age have important implications for water flow through trees. Water supply to the crown is influenced by both the cross-sectional area and the permeability of sapwood. It has been hypothesized that hydraulic conductivity within sapwood increases with age. We investigated changes in sapwood permeability (k) and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans F. Muell. Sapwood was sampled at breast height from trees ranging from 8 to 240 years old, and at three height positions on the main stem of 8-year-old trees. Variation in k was not significant among sampling height positions in young trees. However, k at breast height increased with tree age. This was related to increases in both vessel frequency and vessel diameter, resulting in a greater proportion of sapwood being occupied by vessel lumina. Sapwood hydraulic conductivity (the product of k and sapwood area) also increased with increasing tree age. However, at the stand level, there was a decrease in forest sapwood hydraulic conductivity with increasing stand age, because of a decrease in the number of trees per hectare. Across all ages, there were significant relationships between k and anatomy, with individual anatomical characteristics explaining 33-62% of the variation in k. There was also strong agreement between measured k and permeability predicted by the Hagen-Poiseuille equation. The results support the hypothesis of an increase in sapwood permeability at breast height with age. Further measurements are required to confirm this result at other height positions in older trees. The significance of tree-level changes in sapwood permeability for stand-level water relations is discussed.     

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.     

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

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

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.     

Relationship between the radial variation of ray characteristics and the stages of radial stem increment in Zelkova serrata

We used ten discs from two Zelkova serrata trees - five discs from each tree at sampling heights of 1, 4, 7, 10, and 13 m above the ground - and investigated the radial variation in ray characteristics, i.e., ray area (cross-sectional area of rays on a tangential section), ray density (number of rays/mm² on a tangential section), and ray proportion (the percentage of the area occupied by rays on a tangential section) and analyzed the pattern of variation with respect to the three stages (early, middle, and late) of radial stem increment as estimated using the Gompertz growth function. A juvenile-mature pattern of variation was observed in ray area and ray density. Ray area increased in the inner part of stem and fluctuated around a certain value in the outer part of the stem, and ray density decreased in the inner part of stem and tended to be constant in the outer part of the stem. The maturation age of ray density was similar to the age at the boundary between the early and the middle stages of radial stem increment, but ray area and ray proportion did not relate to the stages of radial stem increment.     

Heartwood and sapwood variation in Eucalyptus globulus Labill. trees at the end of rotation for pulp wood production

The amount of heartwood and sapwood in Eucalyptus globulus Labill. wood was determined in commercial pulpwood plantations at harvest age (12 to 15 years) in four ites in central portugal. twelve trees were sampled in each site at nine stem height levels. Heartwood was present in all the trees up to 82%–87% of the total tree height and amounted to 38%–41% of the total tree volume. The heartwood proportion in the stem cross-section decreased from the base upwards, representing 53.3% and 26.1% of the total area respectively at the base and 55% height level. The sapwood width remained relatively constant along the stem at a mean 38 mm up to the 55% height level. The trees within the site, the height level and their interaction were highly significant sources of variation for heartwood proportion and sapwood width. The heartwood diameter and sapwood width were positively correlated with radial tree growth.     

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

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

Dynamic variation in sapwood specific conductivity in six woody species

Our goals were to quantify how non-embolism-inducing pressure gradients influence trunk sapwood specific conductivity (k(s)) and to compare the impacts of constant and varying pressure gradients on k(s) with KCl and H2O as the perfusion solutions. We studied six woody species (three conifers and three angiosperms) which varied in pit membrane structure, pit size and frequency of axial water transport across pits (long versus short conduits). Both stepwise ("steady") and nonlinear continuous ("non-steady") decreases in the pressure gradient led to decreased k(s) in all species but white oak (Quercus garryana Dougl. ex Hook), a ring-porous and long-vesseled angiosperm. In one diffuse-porous angiosperm (red alder, Alnus rubra Bong.) and two conifers (western red cedar, Thuja plicata Donn. ex D. Don, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco), k(s) was 10-30% higher under steady pressure gradients than under non-steady pressure gradients, and a decrease in the pressure gradient from 0.15 to 0.01 MPa m(-1) caused a 20-42% decrease in k(s). In another diffuse-porous angiosperm (maple, Acer macrophyllum Pursh) and in a third coniferous species (western hemlock, Tsuga heterophylla (Raf.) Sarg), there was no difference between k(s) measured under steady and non-steady pressure gradients. With the exception of western red cedar, a conifer with simple pit membranes, the differences in k(s) between low and high pressure gradients tended to be lower in the conifers than in the diffuse-porous angiosperms. In Douglas-fir, western red cedar and the diffuse-porous angiosperms, k(s) was higher when measured with KCl than with H2O. In white oak, there were no differences in k(s) whether measured under steady or non-steady pressure gradients, or when xylem was perfused with KCl or H2O. The species differences in the behavior of k(s) suggest that elasticity of the pit membrane was the main factor causing k(s) to be disproportionate to the pressure gradient and to the different pressure regimes. The results imply that, if nonlinearities in pressure-flux relationships are ignored when modeling tree water relations in vivo, large errors will result in the predictions of tree water status and its impact on stomatal control of transpiration and photosynthesis.     

Estimating stem maintenance respiration rates of dissimilar balsam fir stands

Stem maintenance respiration rates were measured in five contrasting balsam fir (Abies balsamea (L.) Mill.) stands. At 15 degrees C, average respiration rates for individual stands ranged from 120 to 235 micro mol m(-3) s(-1) when expressed per unit of sapwood volume, from 0.80 to 1.80 micro mol m(-2) s(-1) when expressed per unit of stem surface area, and from 0.50 to 1.00 micro mol g(-1) s(-1) when expressed per unit of nitrogen in the living stem biomass, but differences among stands were not statistically significant. Coefficients of variation ranged from 50 to 100% within stands and were similar for all bases used to express respiration rates. Coefficients of determination for regressions between chamber flux and chamber values of sapwood volume, stem surface area and nitrogen content varied between stands and no one base was consistently higher than the other bases. We conclude that the bases for expressing stem respiration are equally useful. Respiration rates were more closely correlated to stem temperature observed approximately 2 h earlier than to current stem temperature. Among stands, annual stem maintenance respiration per hectare varied from 0.1 to 0.4 Mmol ha(-1) year(-1), primarily because of large differences in sapwood volumes per hectare. Annual stem maintenance respiration per unit of leaf area ranged from 3 to 6 mol m(-2) year(-1), increasing as sapwood volume per hectare increased.     

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

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

Representative heights for assessing whole-tree values of cell-type proportions in Eucalyptus camaldulensis and E. globulus

Eucalyptus camaldulensis Dehnh. and E. globulus Labill. are economically important species for wood and pulpwood materials. Representative heights for assessing whole-tree values of cell-type proportions(vessels, fibers, ray and axial parenchyma percentages) using increment cores were examined by analysis of within-tree variations. Pattern differences were evaluated between trees and species in both radial and axial directions by statistical data analysis(Moses test). In E. camaldulensis,within-tree variation of vessel percentage was generally higher in the upper and outer regions of the trunk. In contrast, E. globulus within-tree variation was unclear. In both species, although no clear pattern of fiber percentages was observed, within-tree variations of ray and axial parenchyma levels were higher in the lower and inner regions.Significant differences in patterns were observed in the axial variation between species for vessel percentages and in the radial variation between trees of E. camaldulensis for ray parenchyma percentages. The representative heights for assessing whole-tree cell-type proportions were 0.8 mabove the ground for E. camaldulensis and 2.8 m for E.globulus, regardless of differences in tree height and pattern of within-tree variation of cell-type proportions.     

Mechanical analysis of the strains generated by water tension in plant stems. Part II: strains in wood and bark and apparent compliance

Tree stems shrink in diameter during the day and swell during the night in response to changes in water tension in the xylem. Stem shrinkage can easily be measured in a nondestructive way, to derive continuous information about tree water status. The relationship between the strain and the change in water tension can be evaluated by empirical calibrations, or can be related to the structure of the plant. A mechanical analysis was performed to make this relationship explicit. The stem is modeled as a cylinder made of multiple layers of tissues, including heartwood, sapwood, and inner and outer bark. The effect of changes in water tension on the apparent strain at the surface of a tissue is quantified as a function of parameters defining stem anatomy and the mechanical properties of the tissues. Various possible applications in the context of tree physiology are suggested.     

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.