Whole-tree water transport scales with sapwood capacitance in tropical forest canopy trees

The present study examines the manner in which several whole‐tree water transport properties scale with species‐specific variation in sapwood water storage capacity. The hypothesis that constraints on relationships between sapwood capacitance and other water relations characteristics lead to predictable scaling relationships between intrinsic capacitance and whole‐tree behaviour was investigated. Samples of sapwood from four tropical forest canopy tree species selected to represent a range of wood density, tree size and architecture, and taxonomic diversity were used to generate moisture release curves in thermocouple psychrometer chambers, from which species‐specific values of sapwood capacitance were calculated. Sapwood capacitance was then used to scale several whole‐tree water transport properties determined from measurements of upper branch and basal sap flow, branch water potential, and axial and radial movement of deuterated water (D₂O) injected into the base of the trunk as a tracer. Sapwood capacitance ranged from 83 to 416 kg m^?3 MPa^?1 among the four species studied and was strongly correlated with minimum branch water potential, soil‐to‐branch hydraulic conductance, daily utilization of stored water, and axial and radial movement of D₂O. The species‐independent scaling of several whole‐tree water transport properties with sapwood capacitance indicated that substantial convergence in plant function at multiple levels of biological organization was revealed by a simple variable related to a biophysical property of water transport tissue.     

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d^–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d^–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m² increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.     

Axial and radial profiles in conductivities, water storage and native embolism in trunks of young and old-growth ponderosa pine trees

Ponderosa pine has very wide sapwood, and yet the spatial and temporal use of that sapwood for water transport is poorly understood. Moreover, there have been few comparisons of function in tips of old-growth trees in comparison with young trees. In the present study, axial and radial specific conductivity (k_s), leaf specific conductivity (LSC), leaf specific conductance (k_l), native embolism and the compartmentalization of sapwood water storage were characterized in trunks of young and old-growth trees. Trunks of young trees had lower k_s, lower LSC and lower native embolism [corresponding to 5% loss of conductivity (PLC)] than trunks of old-growth trees. However, k_l in young trees was 3.5 times higher than in old-growth trees, supporting the hypothesis that tall trees have a reduced ability to transport water to their leaves. Water storage (capacitance) of young trees was not significantly different than at the base of old-growth trees. Although the top of the old-growth trees had similar k_s, LSC and k_l to the young trees for a given cambial age, they had higher native embolism and lower capacitance. There was no trade-off between k_s and native embolism at any height. In the tree crown, outer sapwood had 35–50% higher k_s than the inner sapwood and 17–25 PLC lower native embolism. At the base of the old trees, there was no significant difference in native embolism between the outer, middle and inner sapwood, showing that refilling of embolisms was complete despite the 130-year difference in wood age among these radial positions. Although during the dry season the inner sapwood tended to be more saturated than the outer sapwood, the outer part of the sapwood contributed up to 60% of the overall stored water. Safer xylem, higher capacitance and higher k_l would appear adaptive in the young trees for regulating their water resource, which is likely to be less reliable than the water availability of older trees with their more developed root system.     

Dynamics of water transport and storage in conifers studied with deuterium and heat tracing techniques

The volume and complexity of their vascular systems make the dynamics of long-distance water transport in large trees difficult to study. We used heat and deuterated water (D2)) as tracers to characterize whole-tree water transport and storage properties in individual trees belonging to the coniferous species Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. The trees used in this study spanned a broad range of height (13.5-58 m) and diameter (0.14-1.43 m). Sap flow was monitored continuously with heat dissipation probes near the base of the trunk prior to, during and following injection of D2O. The transit time for D2O transport from the base of the trunk to the upper crown and the tracer residence time were determined by measuring hydrogen isotope ratios in water extracted from leaves sampled at regular intervals. Transit times for arrival of D2O in the upper crown ranged from 2.5 to 21 d and residence times ranged from 36 to 79 d. Estimates of maximum sap velocity derived from tracer transit times and path length ranged from 2.4 to 5.4 m d(-1). Tracer residence time and half-life increased as tree diameter increased, independent of species. Species-independent scaling of tracer velocity with sapwood-specific conductivity was also observed. When data from this study were combined with similar data from an earlier study of four tropical angiosperm trees, species-independent scaling of tracer velocity and residence time with sapwood hydraulic capacitance was observed. Sapwood capacitance is an intrinsic tissue-level property that appears to govern whole-tree water transport in a similar manner among both tracheid- and vessel-bearing species.     

Sapflow+: a four-needle heat-pulse sap flow sensor enabling nonempirical sap flux density and water content measurements

? To our knowledge, to date, no nonempirical method exists to measure reverse, low or high sap flux density. Moreover, existing sap flow methods require destructive wood core measurements to determine sapwood water content, necessary to convert heat velocity to sap flux density, not only damaging the tree, but also neglecting seasonal variability in sapwood water content. ? Here, we present a nonempirical heat-pulse-based method and coupled sensor which measure temperature changes around a linear heater in both axial and tangential directions after application of a heat pulse. By fitting the correct heat conduction-convection equation to the measured temperature profiles, the heat velocity and water content of the sapwood can be determined. ? An identifiability analysis and validation tests on artificial and real stem segments of European beech (Fagus sylvatica L.) confirm the applicability of the method, leading to accurate determinations of heat velocity, water content and hence sap flux density. ? The proposed method enables sap flux density measurements to be made across the entire natural occurring sap flux density range of woody plants. Moreover, the water content during low flows can be determined accurately, enabling a correct conversion from heat velocity to sap flux density without destructive core measurements.     

云南哀牢山6种常绿阔叶树木质部解剖特征的轴向和径向变化

West、Brown和Enquist提出的树木水分传导的分形网络模型(简称WBE模型)认为,树木连续分枝之间的导管或管胞直径按照一定的比率均匀变细,其总的水力阻力与水分传导的路径长度无关,从而使不同部位叶片获得基本相当的水分供应。该模型对树木高生长的水力限制假说提出了置疑。为了验证WBE模型中树木导管或管胞均匀变细的假说,该文研究了云南哀牢山中湿性常绿阔叶林中6种常绿阔叶树, 腾冲栲(Castanopsis wattii)、景东石砾(Lithocarpus chintungensis)、木果石砾(L. xylocarpus)、长尾青冈(Cyclobalanopsis stewardiana)、滇木荷(Schima noronhae)和舟柄茶(Hartia sinensis)木质部解剖特征随树高和年龄的变化。对这6个树种共14株样木进行了不同高度树干圆盘和边材生长轮取样,样木的高度为15~25 m,按照常规木材解剖的处理和分析方法,在显微镜下测定木材切片的导管直径和密度等特征。结果表明:在14株样木中,有4株树木导管直径随树木高度增加呈线性减小, 1株没有明显变化,其它9株树木导管直径在树冠以下的树干部分变化幅度较小或没有明显变化,而从树冠基部往上直到树木顶端导管直径显著减小。同一植株随着高度的增加,导管密度增加并且在树冠内增加更显著。有三分之一的树木导管占边材面积的比例随树高增加没有明显变化,其余树木导管占边材面积比在树冠以上有所减小。多数树木理论比导率在树冠以下没有明显变化而在树冠基部往上显著降低。在从髓芯开始往外的20~40个年轮范围内导管直径增加显著,但大部分植株导管直径在40个年轮后趋于稳定。不同高度圆盘导管直径随形成层发育时间的变化呈相似的趋势,并且相同发育年龄的导管直径没有明显差异。该文的研究结果说明,导管直径的轴向和径向变化一定程度上补偿了水分运输阻力随树木个体增大而增加的缺陷,但是6种常绿阔叶树树干的导管基本不按一定比率均匀变细,不支持WBE模型。     

Radial patterns of xylem sap flow in non-, diffuse- and ring-porous tree species

We investigated radial patterns of sap flux density and wood properties in the sapwood of young loblolly pine (Finns taeda L.), mature white oak (Quercus alba L.) and sweetgum (Liquidambar styraciflua L.), which represent three major classes of wood anatomy: non-porous (coniferous), ring-porous and diffuse-porous. Radial measurements of xylem sap flux density were made in sections of xylem extending to 20 mm and 20–40 mm from the cambium. These measurements were compared with measurements of the relative water content (R_s) and sapwood specific gravity (ρ_r) of corresponding radial sections. In both hardwood species, sap flow differences were rarely significant between the two depth intervals. In pine, a 59% reduction in daily sap flux density from outer to inner sapwood was found. This could not be accounted for by a 3% drop in R_s; rather, an accompanying 9% reduction in ρ_r indicated a transition between the depth intervals from mature to juvenile sapwood, and is the probable cause of the lower flux rate in the inner xylem of pine.     

A novel approach to the in situ measurement of oxygen concentrations in the sapwood of woody plants

A novel technique for the physico‐chemical analysis of xylem sap by underwater access to the sapwood of trees is described. In situ measurements of dissolved oxygen in the sapwood are performed by combining this technique with a novel optical method for oxygen detection. In early spring, the oxygen concentration of the sapwood of Betula pendula was in the range of 80–230 µmol O₂ L^?1, corresponding to an oxygen deficit of 40–75% of air saturation. Oxygen concentration maxima and minima occurred early in the morning and in the afternoon, respectively, whereas xylem sap temperatures showed the reverse pattern. In the sapwood, hypoxia increased from the beginning of bud break until frondescence, when a deficit of 86% of air saturation marked the upper limit of oxygen depletion. There seemed to be no relationship between daily variations of oxygen concentration and xylem sap pressure. In summer, sap flow was a major determinant for the diurnal variation of dissolved oxygen concentration. Oxygen supply to the sapwood was determined by both radial influx into the trunk through intercellular gas spaces and transport of dissolved oxygen via xylem sap flow. Radial influx seemed to be favoured during night‐time, when the trunk was warmer than ambient air. During daytime, the hypoxia of the sapwood rose and increased sharply in the evening, when sap flow velocity approximated zero. High temperature in the sapwood enhanced the respiratory oxygen consumption of the wood parenchyma while the supply of dissolved oxygen via the transpiration stream became ineffective.     

Granier树干液流测定系统在马占相思的水分利用研究中的应用

介绍了Granier热消散探针在树干液流测定中的工作原理,并利用该系统长期监测广东鹤山马占相思林14株样树的液流密度,分析了树木个体内和个体之间液流密度的差异、整树和林段水分利用的量化特征.由于树木边材结构以及周围微环境的差别,树木内和个体间的液流密度差异非常明显,变异系数的平均值分别为15.51%-37.26%、37.46%-50.73%.尽管液流密度的差异较大,但同一株树木不同方位的液流密度之间却呈现明显的线性相关（p＜0.0001）,这是重要的特征值,使得只需测定某一方位的液流密度经尺度外推计算整树和林段蒸腾成为可能.树木液流对环境因子响应的变化规律取决于所参照的时间尺度,日变化主要受光辐射、水汽压差等气候因子的控制,而土壤水份对液流的季节变化影响较大.形态特征明显影响树木的液流,高大树木由于边材较厚、树干粗壮和冠幅较宽而承载较多的辐射能量,因而水分蒸腾较高.对树木液流密度在径向和方位上进行适当的整合,可较准确地计算整树和林段蒸腾.由液流估测的马占相思整树和林段蒸腾的结果显示,该群落的水分利用在时间和空间上均有明显的分化.     

Climate-related trends in sapwood biophysical properties in two conifers: avoidance of hydraulic dysfunction through coordinated adjustments in xylem efficiency, safety and capacitance

In the Pacific north‐west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas‐fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas‐fir. Radial conductivity was higher in west‐side populations of both species, but axial conductivity was higher in the east‐side populations and in Douglas‐fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west‐side populations. Sapwood capacitance was similar between species, but was about twice as great in east‐side populations (580 kg m^?3 MPa^?1) as in west‐side populations (274 kg m^?3 MPa^?1). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity.     

Scots pine root distribution derived from radial sap flow patterns in stems of large leaning trees

This study characterizes whole tree root system distribution in a non-destructive way based on its functional parameters, particularly the sap flow patterns in stems. This approach particularly considers sap flow variation across stems, both radial and circumferential patterns of flow that are usually used for a better integration of sap flow density at the whole tree level. We focused at: (1) Showing examples of sap flow variation across stems at a defined situation (high midday values at the period of non-limiting water supply; (2) Analyzing radial flow patterns in terms of root distribution; (3) Validating these results at the stand level (mean data of series of individual trees) using results of classical biometric methods used at the same site; and (4) Applying the results for evaluation of root distribution around leaning trees. Sap flow rate was measured by the heat deformation method on a set of 14 trees at an experimental pine forest stand in Brasschaat (Belgium) during the growing season of 2000. Sap flow variation across stems was measured at a total of 700 points. Amounts of water supplied by superficial (horizontally oriented) and sinker (vertically oriented) roots were estimated from sap flow patterns. The vertical distribution of absorbing roots as derived from the analysis of sap flow patterns in stem sapwood was very similar to the distribution determined by the classical biometric analysis of fine roots. Trees leaning to the East had stem radii at the stump level and crown radii enhanced in the leaning direction. Sinker roots showed higher absorption activities in the leaning direction, but superficial roots were more absorbing in the opposite direction. The application of the above-described method allows for a better evaluation of the whole-tree behavior and facilitates the evaluation of tree and stand properties in traditional forest stands, which are not equipped for detailed scientific research. This may also facilitate practical applications in landscape-level studies.     

Maple sap uptake, exudation, and pressure changes correlated with freezing exotherms and thawing endotherms

Sap flow rates and sap pressure changes were measured in dormant sugar maple trees (Acer saccharum Marsh.). In the forest, sap flow rates and pressure changes were measured from tap holes drilled into tree trunks in mature trees and sap flow rates were measured from the base of excised branches. Excised branches were also brought into the laboratory where air temperature could be carefully controlled in a refrigerated box and sap flow rates and sap pressures were measured from the cut base of the branches.

Under both forest and laboratory conditions, sap uptake occurred as the wood temperature declined but much more rapid sap uptake correlated with the onset of the freezing exotherm. When sap pressures were measured under conditions of negligible volume displacement, the sap pressure rapidly fell to −60 to −80 kilopascals at the start of the freezing exotherm. The volume of water uptake and the rate of uptake depended on the rate of freezing. A slow freezing rate correlated with a large volume of water uptake, a fast freezing rate induced a smaller volume of water uptake. The volume of water uptake ranged from 0.02 to 0.055 grams water per gram dry weight of sapwood. The volume of water exuded after thawing was usually less than the volume of uptake so that after several freezing and thawing cycles the sapwood water content increased from 0.7 to 0.8 grams water per gram dry weight.

These results are discussed in terms of a physical model of the mechanism of maple sap uptake and exudation first proposed by P. E. R. O'Malley. The proposed mechanism of sap uptake is by vapor distillation in air filled wood fiber lumina during the freezing of minor branches. Gravity and pressurized air bubbles (compressed during freezing) cause sap flow from the canopy down the tree after the thaw.

     

Crown conductance in dwarf, medium, and tall pitch pines in the Long Island Pine Plains

The New York Pine Plains are a unique ecosystem with normal statured and a dwarfed variety of pitch pines (Pinus rigida Mill.). Growing interspersed with the dwarf pines are trees of intermediate height and features. Several hypotheses have been put forward as to why some of the trees are dwarfed, but none have been substantiated. In this study, we tested whether dwarf or medium trees are hydraulically limited compared to normally growing trees. Granier style sap flux sensors were installed in three to six trees of each tree type and sap flux was measured in early August 2004. Sap flux measurements were scaled to crown stomatal conductance using leaf area to sapwood area ratios for each tree. Contrary to expectation, dwarf and medium stature trees had very low leaf area to sapwood area ratios, but high crown stomatal conductances compared to normal trees. Analyses of leaf area, ring widths, and crown stomatal conductance indicate that differences between normal, and dwarf and medium pines are not due to hydraulic limitation, but that stunted growth may be due to other causes.     

Axial Water Flux Dynamics in Small Diameter Roots of a Fast Growing Tropical Tree

Field measurements of water flux in small diameter roots are important to the study of whole plant water transport systems. Miniature sap flow gauges were used to capture high resolution water flux patterns in small roots of 2 – 5 mm diameter and simultaneously in the canopy branches of a Eucalyptus saligna tree growing in Hawaii. The axial transport flux rates were then correlated with anatomical measurements to describe the internal hydraulic capacity of the tree. The daily patterns of water flux showed a strong coupling between the canopy and root systems and both systems were tightly synchronized with rapid fluctuations in photosynthetic photon flux density, vapour pressure deficit, and wind speed. When flow rates were normalized by the total vessel lumen area, branches had daily totals equivalent to the surface roots. Daily flows of water through surface roots were consistently 30% greater than through deep roots. Results of an experiment where a portion of the canopy was removed showed the decrease in water flux for all roots was in nearly direct proportion to the decrease in leaf area. The root anatomical measurements suggested a high capacity axial root water transport system with roots containing a smaller number of vessels per unit of sapwood area than branches but with vessel diameters twice that of the branches. However, relative conductivity values of roots and branches were similar and comparable to some of the highest values reported. Overall, the results suggested a highly efficient axial water transport system that would help to maintain a favorable plant water status for maximal stomatal opening. This revised version was published online in July 2006 with corrections to the Cover Date.     

Tree water dynamics non-destructively assessed through sap flow measurements and potential evapotranspiration

Sap flow and potential evapotranspiration rates were analyzed for two coniferous tree species (Douglas-fir and Scots pine) and one broadleaf species (sessile oak) in a mixed Carpineto-Quercetum forest during the growing season 2005. The relationship between sap flow and potential evapotranspiration rates, effective crown area as a measure of the relative transpiration and daily relative proportion of the storage water used for transpiration were used as indicators of the tree water dynamics. These indicators were determined on four consecutive days and all three showed good reliability concerning tree water dynamics.     

桉树人工林树液流动密度随边材径向深度的变化

树液流动密度 (SFD)随边材径向深度的变化对于准确估测流经边材的树液通量是非常重要的 ,后者又制约着HeatPulse的应用精度。但迄今为止 ,只有很少的研究估计了由于SFD随径向的梯度变化而带来的误差 ,SFD沿树干径向分布规律的获得往往依靠对少数几棵树的观测资料。基于在广东雷州半岛对两块 3～ 4年生桉树 (Euca lyptusurophyllaS .T .Blake)人工林 1年的HeatPulse观测 ,探讨了对来自 39株立木大量观测资料的综合处理方法 ,发现这两个样地 (纪家和河头 )的林分中SFD随边材径向深度的变化可以用如下回归方程来描述 :纪家 :y =3.6 6 75x3 - 7.2 95 5x2 3.6 82 6x 0 .5 6 74 (R2 =0 .9391,n =80 ,P =0 .0 1)河头 :y =5 .0 0 6 2x3 - 9.116 1x2 4.4 5 4 4x 0 .4 6 34(R2 =0 .80 6 9,n =72 ,P =0 .0 1)式中 :y———某一树液感应器所测得的SFD与不同深度的 4个感应器所测得的SFD的平均值之比 ;x—某一树液感应器在边材中的深度与边材厚度之比。从形成层到心材 ,SFD最初有所增加 ,随后持续减小 ,但由于树木年龄很小 ,最大的SFD只比最小的SFD大 0 .33～ 0 .36倍。     

Evaluation of the heat pulse velocity method for measuring sap flow in Pinus patula

Information on the water use of Pinus patula plantations isrequired to predict the impact of forest plantations on waterresources in South Africa. The heat pulse velocity (HPV) methodis a promising technique for measuring water use by trees, andhas been shown to measure sap flows accurately in a varietyof hardwood trees. This method has not been sufficiently verifiedfor pine trees where the presence of a strongly-defined ringstructure in the sapwood gives rise to a complex radial patternof sap flow. The purpose of this study was to compare wateruptake by cut trees to simultaneous HPV sap flow measurementsin the same tree. Fourteen trees were used for this comparison.Results showed that HPV sap flow estimates consistently overestimatedcut-tree uptake by an average of 49%. The bias is attributedto heat averaging across non-conducting latewood rings. Wateruptake was found to be highly correlated to the product of under-barkcross-sectional area and wound-corrected mean HPV, and it issuggested that this empirical relation provides a more appropriateway of estimating water use by this species. Key words: Heat pulse velocity, sap flow, Pinus patula, transpiration     

Xylem sap flow as a major pathway for oxygen supply to the sapwood of birch (Betula pubescens Ehr.)

The role of xylem sap flow as an aqueous pathway for oxygen supply to the wood parenchyma of Betula pubescens saplings was investigated. Using micro‐optode sensors the oxygen status of the sapwood was quantified in relation to mass flow of xylem sap. Sap flow was gradually reduced by an increasing oxygen depletion in the root space. The effect of sap flow on radial O₂ transport between stem and atmosphere was assessed by a stoichiometrical approach between respiratory CO₂ production and O₂ consumption. Restriction of sap flow set in 36.5 h after the onset of O₂ depletion, and was complete after 71 h. Interruption of sap flow drastically increased the O₂ deficit in the sapwood to 70%. Sap flow contributed about 60% to the total oxygen supply to the sapwood. Diurnal O₂ flow rates varied between 3 and 6.3 nmol O₂ m^?2 leaf area (LA) s^?1 during night‐ and daytime, respectively. Maximum O₂ flow rates of 20 nmol O₂ m^?2 LA s^?1 were reached at highest sap flow rates of 5.7 mmol H₂O m^?2 LA s^?1. Sap flow not only affected the oxygen status of the sapwood but also had an effect on radial O₂ transport between stem and atmosphere.     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

Use of the correct heat conduction-convection equation as basis for heat-pulse sap flow methods in anisotropic wood

Heat-pulse methods to determine sap flux density in trees are founded on the theory of heat conduction and heat convection in an isotropic medium. However, sapwood is clearly anisotropic, implying a difference in thermal conductivity along and across the grain, and hence necessitates the theory for an anisotropic medium. This difference in thermal conductivities, which can be up to 50%, is, however, not taken into account in the key equation leading to the currently available heat-pulse methods. Despite this major flaw, the methods remain theoretically correct as they are based on derivations of the key equation, ruling out any anisotropic aspects. The importance of specifying the thermal characteristics of the sapwood according to axial, tangential or radial direction is revealed as well as referring to and using the proper anisotropic theory in order to avoid confusion and misinterpretation of thermal properties when dealing with sap flux density measurements or erroneous results when modelling heat transport in sapwood.