Effects of tension wood on specific conductivity and vulnerability to embolism of Quercus ilex seedlings grown at two atmospheric CO2 concentrations

To determine whether there are decreases in hydraulic function of a woody stem when it has increased mechanical loading, Quercus ilex L. seedlings were grown upright or inclined to force the production of large amounts of tension wood (TW). Seedlings were grown in ambient or elevated carbon dioxide concentrations ([CO2]) for 16-17 months to provide two sets of seedlings differing in growth rates and allocation patterns. In both CO2 environments, inclination caused formation of large amounts of TW at the base and mid-section of most stems, but not at the stem tips. Contrary to expectation, there were no significant effects of stem inclination or amount of TW on specific conductivity (k(s)) or vulnerability to embolism. Samples with high amounts of TW had higher vessel frequency, similar average vessel lumen area, similar vessel lumen fraction (6% of the transverse area), elevated frequency of vessels in the smallest diameter class, and higher wood density than samples with very little TW. Samples from seedlings in the elevated [CO2] treatment had similar vessel frequency, larger average vessel lumen area (caused by a higher frequency of large-diameter vessels), similar vessel lumen fraction, and similar wood density as samples from seedlings in the ambient [CO2] treatment. There was a strong position effect: the highest wood density and lowest ks were at the stem base, intermediate values were at the middle, and the lowest density and highest ks were at the stem tip. We conclude that, in a species that uses different cells for mechanical support and water transport, there can be large modifications in performance of the mechanical function through TW formation without impacting the water transport functions-ks and vulnerability to embolism.     

Wood density and anatomy of water-limited eucalypts

We hypothesized that seedlings grown under water-limited conditions would develop denser wood than seedlings grown under well-watered conditions. Three Eucalyptus species (E. grandis Hill (ex Maiden), E. sideroxylon Cunn. (ex Woolls) and E. occidentalis Endl.) were grown in a temperature-controlled greenhouse for 19 weeks with watering treatments (well-watered and water-limited) applied at six weeks. The water-limitation treatment consisted of four drought cycles. Wood density increased by between 4 and 13% in the water-limited seedlings, but this increase was mainly due to extractive compounds embedded in the cell wall matrix. Once these compounds were removed, the increase was 0-9% and was significant for E. grandis only. Water-limitation significantly reduced mean vessel lumen area; however, this was balanced by a trend toward greater vessel frequency in water-limited plants, and consequently there was no difference in the proportion of stem area allocated to vessels. Conduit efficiency value was lowest in the water-limited plants, indicating that there was a cost in terms of stem hydraulic conductivity for decreasing vessel lumen area. Wood density was negatively correlated with vessel lumen fraction in well-watered plants, but this relationship broke down in the water-limited plants, possibly because of the significantly larger proportion of the stem taken up by pith in water-limited seedlings. Diurnal variation in leaf water potential was positively correlated with wood density in well-watered plants. This relationship did not hold in the water-limited plants owing to the collapse of the pressure gradient between soil and leaf. We conclude that drought periods of greater than 1 month are required to increase wood density in these species and that increases in wood density appear to result in diminished capacity to supply water to leaves.     

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.     

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.     

Does growing on a slope affect tree xylem structure and water relations?

Variations in slope, exposure, relief and substrate over a short distance and their influences on plant function are poorly understood. We investigated the influences of soil hydrological characteristics on internal stem structure and hydraulic properties of downy oak (Quercus pubescens Willd.) growing along a hill slope. Increment wood cores were extracted from the base and at breast height (BH) of tree stems. Relative wood water content (W(c)) and wood density (D(w)) were measured in the sapwood. Wood compression strength (delta) in the longitudinal direction was measured with a fractometer. Thin sections were cut from the transversal face of each core and vessel lumen area (V(A)) was measured and xylem theoretical hydraulic conductivity (L(th)) estimated over the sapwood. Topsoil volumetric water content (theta(v)) was determined around trees and the hydrological behavior of the slope was studied through field surveys. Data were used as input to a hydrological model to simulate topsoil water distribution along the slope. Results showed that theta(v) tends to decrease with increasing altitude. Groundwater levels were lower upslope than downslope, and results from the hydrological model confirmed these trends. Mean W(c) at the base of each tree decreased significantly with increasing altitude, whereas at BH, no differences were found along the slope. There was a significant positive relationship between W(c) measured at the tree base and theta(v) along the hill slope, but not for W(c) measured at BH. Values of D(w) and delta measured at both stem positions increased significantly with increasing altitude and decreasing theta(v). Significant negative relationships were found between delta and theta(v) measured at the stem base and at BH. At both stem positions, delta was closely related to D(w) and L(th). Vessel lumen areas at BH and the stem base were significantly regressed with altitude, theta(v), D(w) and delta. Xylem theoretical hydraulic conductivity at both stem positions was negatively related to altitude and soil theta(v), but only L(th) measured at the stem base was negatively regressed with D(w). The results are discussed in the context of how tree position along a hill slope influences water uptake and internal xylem structure.     

Effects of ring-porous and diffuse-porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model

Calibration of a recently developed water flow and storage model based on experimental data for a young diffuse-porous beech tree (Fagus sylvatica L.) and a young ring-porous oak tree (Quercus robur L.) revealed that differences in stem wood anatomy between species strongly affect the calibrated values of the hydraulic model parameters. The hydraulic capacitance (C) of the stem storage tissue was higher in oak than in beech (939.8 versus 212.3 mg MPa(-1)). Model simulation of the elastic modulus (epsilon) revealed that this difference was linked to the higher elasticity of the stem storage tissue of oak compared with beech. Furthermore, the hydraulic resistance (R (x)) of beech was about twice that of oak (0.1829 versus 0.1072 MPa s mg(-1)). To determine the physiological meaning of the R (x) parameter identified by model calibration, we analyzed the stem wood anatomy of the beech and oak trees. Calculation of stem specific hydraulic conductivity (k (s)) of beech and oak with the Hagen-Poiseuille equation confirmed the differences in R (x) predicted by the model. The contributions of different vessel diameter classes to the total hydraulic conductivity of the xylem were calculated. As expected, the few big vessels contributed much more to total conductivity than the many small vessels. Compared with beech, the larger vessels of oak resulted in a higher k (s) (10.66 versus 4.90 kg m(-1) s(-1) MPa(-1)). The calculated ratio of k (s) of oak to beech was 2, confirming the R (x) ratio obtained by model calibration. Thus, validation of the R (x) parameter of the model led to identification of its physiological meaning.     

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.     

木材导热系数的研究

本文运用非平衡态热力学线性理论分析了含水率在纤维饱和点以下木材的传热,结果表明木材中的水蒸汽和束缚水参与了传热,木材导热系数由传导导热系数与因水蒸汽和束缚水扩散产生的两个等效导热系数组成。推广Siau关于木材细胞的导热理论,依据水分吸附和扩散的原理,导出了木材弦向和径向导热系数计算公式,与已发表的36种气干材弦向实验值和23种气干材径向实验值符合较好。分析木材径向传热的方法也适用于木材纵向传热。     

The anatomical traits of trunk wood and their relevance to oak (Quercus robur L.) vitality

Oaks’ decline in vitality is attributed to a complex process that involves interactions of several factors leading to increased trees’ mortality. This study investigates the structure of trunk wood of oaks with reference to its physiological role in hydraulic conductivity. On the basis of the crown condition, the oaks were classified into three health groups: healthy trees, declining trees and dead trees. Anatomical traits of wood, such as annual ring width, vessel density, vessel diameter of earlywood and theoretical hydraulic conductivity, were measured and calculated. The narrowest annual rings formed by the cambium were observed in dead oaks. These trees were also characterized by the smallest diameter of earlywood vessels, not only in the period of occurrence of dieback symptoms, but also during their whole life. It is suggested that the formation of narrow annual rings and earlywood vessels of small diameter increases susceptibility of a tree to decay. A reduced vessel diameter implies changes in hydraulic conductivity of oak trunks and thus impairs the water transport, which affects the health of trees. The process of oak decline is considered to have characteristics of natural selection and leads to the elimination of the weakest trees.     

Refined pipe theory for mechanistic modeling of wood development

We present a mechanistic model of wood tissue development in response to changes in competition, management and climate. The model is based on a refinement of the pipe theory, where the constant ratio between sapwood and leaf area (pipe theory) is replaced by a ratio between pipe conductivity and leaf area. Simulated pipe conductivity changes with age, stand density and climate in response to changes in allocation or pipe radius, or both. The central equation of the model, which calculates the ratio of carbon (C) allocated to leaves and pipes, can be parameterized to describe the contrasting stem conductivity behavior of different tree species: from constant stem conductivity (functional homeostasis hypothesis) to height-related reduction in stem conductivity with age (hydraulic limitation hypothesis). The model simulates the daily growth of pipes (vessels or tracheids), fibers and parenchyma as well as vessel size and simulates the wood density profile and the earlywood to latewood ratio from these data. Initial runs indicate the model yields realistic seasonal changes in pipe radius (decreasing pipe radius from spring to autumn) and wood density, as well as realistic differences associated with the competitive status of trees (denser wood in suppressed trees).     

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.     

Selecting earlywood vessels to maximize their environmental signal

The anatomical features of earlywood vessels often reflect information about past climatic conditions. We examined the relationships between mean monthly temperature and mean vessel lumen area (MVA) in various categories of earlywood vessels. Subsets of earlywood vessels of chestnut (Castanea sativa Mill.) were selected from a previously reported dataset based on several progressive size-related procedures. To include all earlywood vessels, the minimum size considered was 10,000 microm2. Changes in the correlations between MVA and the mean air temperature in March are described and discussed. The results show that not all vessels embody the same information. The MVA of a proportion of the largest earlywood vessels in each annual ring was most closely related to March temperature, whereas MVA of the smallest earlywood vessels was better correlated with June temperature. This difference is probably a result of the vessels being formed at different times: early spring for the largest earlywood vessels and later in spring for the smallest earlywood vessels. Analyses combining large and small vessels yielded lower correlations between MVA and monthly temperature. The number, size and distribution of vessels can vary greatly from ring to ring. In making year-to-year comparisons, the best information is provided by observations on vessels of contemporaneous ontogenesis. Criteria for the selection and analysis of vessels in the assessment of temperature during the season of wood formation are proposed and discussed.     

木麻黄树干细胞组织结构与对星天牛抗虫性的关系

本文报道了４个品系木麻黄木材组织显微解剖结果，分析了木材导管、导管间区木化细胞和射线等物理特征，探讨了木麻黄抗虫机制。高抗品系Ｃ４４木材导管数量少，单位面积（１ｍｍ２）仅有２０．２５个导管，且占木材面积少，仅为９．７２％；导管间区木化细胞较明显，壁厚与腔径比值最大，为５６．５９；单位面积射线的束数、长度、宽度和射线面积／总面积等指标均为最低。说明该品系木麻黄木质部水分含量低、材质坚硬致密、木质部提供养分少，因而不利于星天牛幼虫的生存发育。     

Diminished vessel diameter as a possible factor in the decline of European ash (Fraxinus excelsior L.)

? The aim of this work was to examine the anatomy and functioning of secondary xylem in stems in relation to the decline of European ash (Fraxinus excelsior L.).

? We tested the hypothesis that declining trees show changes in the structure of wood, which result in impaired water transport.

? Anatomical analyses were carried out on wood samples (comprising all annual rings formed during the 30 years life of the analysed trees) collected at breast height from the main stem of healthy, weakened and dead ash trees. The width of annual wood increments, the diameter and density of earlywood vessels were measured and the theoretical hydraulic conductivity index through the secondary xylem calculated by application of Hagen-Poisseuille’s formula.

? Anatomical characteristics changed both with the age of trees and in response to unfavorable factors. The largest vessels were observed in healthy trees, which implied that they had the highest hydraulic conductivity index, whereas trees considered to be in decline produce smaller vessels and hence had reduced conductivity.

     

木材横纹导热系数的类比法研究

分析导热与导电间某些属性的相似性 ,应用类比法推出导热系数具有导电系数的某些属性 ,可用导电系数类似的定义式定义导热系数 ,即木材导热系数等于木材单位长度单位截面的热阻的倒数。根据木材微观细胞形态 ,选用圆柱形模型推导木材横纹导热系数的理论计算式。应用该公式计算 2 0种木材的横纹导热系数 ,理论值的最大误差 14 1% ,平均误差 7% ,理论值与试验值较为吻合 ,为理论研究木材热学性质提供一种可适用的方法。     

Limitations of a compensation heat pulse velocity system at low sap flow: implications for measurements at night and in shaded trees

Unlike an ideal system, the return time to thermal balance (t(b)) between upstream and downstream thermistors, as measured by the (compensation) heat pulse velocity method, effectively depends on the heat input and the water content of the wood at zero and low sap flow. Even when these factors were held constant and ambient temperature was stabilized, a twofold variation in t(b) at zero flow was observed within and among Greenspan Technology sensors implanted in wooden posts, making it impossible to distinguish zero flow from low sap velocities (< 0.01-0.02 mm s(-1)). This limitation has serious consequences because the contribution of low flow rates to water movement is important during both daytime and nighttime in tropical understory and overstory trees. Measurements in an artificial flow system showed that this technical limitation is exacerbated by erratic variation in sensor response at both zero and low flow rates. The limited sensitivity of the tested sap flow sensors may be caused by their poor thermal contact with wood. Interim procedures are suggested for estimating minimum detectable sap flow and delimiting the hydroactive zone until the sensitivity and interchangeability of sap flow probes are improved.     

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.     

Hydraulic conductivity, photosynthesis and leaf water balance in six evergreen woody species from fall to winter

To confirm that freeze-thaw embolism is a primary stress for evergreen woody species in winter, hydraulic conductivity, photosynthesis and leaf water potential were measured during fall and winter in trees growing in a cool temperate zone (Nikko) and in a warm temperate zone (Tokyo). We examined two evergreen conifers that naturally occur in the cool temperate zone (Abies firma Siebold & Zucc. and Abies homolepis Siebold & Zucc.), and four evergreen broad-leaved woody species that are restricted to the warm temperate zone (Camellia japonica L., Cinnamomum camphora (L.) J. Presl, Ilex crenata Thunb. and Quercus myrsinaefolia Blume). In Tokyo, where no freeze-thaw cycles of xylem sap occurred, hydraulic conductivity, photosynthesis and water balance remained constant during the experimental period. In Nikko, where there were 38 daily freeze-thaw cycles by February, neither of the tracheid-bearing evergreen conifers showed xylem embolism or leaf water deficits. Similarly, the broad-leaved evergreen trees with small-diameter vessels did not exhibit severe embolism or water deficits and maintained CO(2) assimilation even in January. In contrast, the two broad-leaved evergreen trees with large-diameter vessels showed significantly reduced hydraulic conductivity and shoot die-back in winter. We conclude that freeze-thaw embolism restricts evergreen woody species with large-diameter vessels to the warm temperate zone, whereas other stresses limit the distribution of broad-leaved trees, that have small-diameter vessels, but which are restricted to the warm temperate zone.     

Systematic variation in xylem hydraulic capacity within the crown of white ash (Fraxinus americana)

A 7-m tall white ash tree (Fraxinus americana Marsh.) was dissected, and hydraulic parameters of the xylem were determined by inducing a steady-state flow of water through the stem segments and monitoring volume and velocity flow rates. Leaf-specific conductivity (LSC) was highest in the main stem and lowest in some of the leaf-bearing lateral shoots. The LSC was higher in the main stem than in branches and higher in primary than in secondary branch axes. Terminal leaf-bearing shoots were larger and had a significantly greater mean LSC than subjacent lateral shoots. A significant reduction in LSC was associated with the transition between 1- and 2-year-old growth. In branches of the same age, there was a close correspondence among LSC, branch position and branch size. The average LSC of leaf-bearing shoots from south-facing branches was 43% greater than that of shoots from north-facing branches. Within-crown variation in LSC was associated with variation in velocity flow rate (V). By contrast, the ratio of potentially functional xylem area to supported leaf area (A(pf)/A(l)) was relatively stable throughout the crown. Stratification of stems by Strahler order accounted for approximately 70% of the total variation in LSC. These results suggest that (1) there exists a systematic pattern of variation in LSC distribution within the crown of white ash, (2) within-crown variability in LSC is primarily the result of variation in mean vessel diameter, and (3) there is a physiological linkage between LSC and crown morphology that is maintained through a positive feedback mechanism during branch ontogeny.     

Climatic signal of earlywood vessels of oak on a maritime site

Earlywood vessel lumen areas were measured in 72 consecutive tree rings in wood cores from oak (Quercus robur L.) trees in a maritime woodland. This anatomical time series was statistically correlated with climate data for the same time span. There was a strong dependence of earlywood vessel lumen area on rainfall between February and April, which reflects the role of water availability in vessel ontogeny. By inversion, earlywood vessel lumen areas can be used as a proxy to reconstruct spring precipitation beyond the archived weather records. Such information may be of value in the context of climate change.