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.     

Evaluation of the heat pulse velocity technique for measurement of sap flow in rainforest and eucalypt forest species of south-eastern Australia

Sap flow in the stems of two cut saplings each of Eucalyptus maculata (a canopy eucalypt forest tree), Doryphora sassafras and Ceratopetalum apetalum (both canopy rainforest trees of south-eastern coastal Australia) was measured by the heat pulse velocity technique and compared with water uptake from a potometer. Scanning electron micrographs of wounding caused by implantation of temperature sensor and heater probes into the sapwood showed that wounding was similar in rainforest and eucalypt species and was elliptical in shape. A circular wound has been implicitly assumed in previous studies. Accurate measurements of sapling water use were obtained using the smaller transverse wound dimension rather than the larger longitudinal dimension because maximum disruption of sap flow through the xylem vessels occurred in the transverse plane. Accurate measurements of sap flux were obtained above a minimum threshold sap velocity. These velocities were 15·7,10·9 and 9·4 cm h^?1 for E. maculata, C. apetalum and D. sassafras, respectively. Below the threshold sap velocity, however, sap flow could not be accurately calculated from measurements of heat pulse velocity. The minimum threshold sap velocity appeared to be determined by probe construction and xylem anatomy. Despite the elliptical wounding and inaccurate measurement of sap flow below the threshold sap velocity, total sap flow over the experimental period for two saplings of each species was within 7% of water use measured by the potometer.     

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.     

Measurement of sap flow in plant stems

Transpiration rates for whole plants, individual branches ortillers can be determined by techniques which measure the rateat which sap ascends stems. All of these methods use heat asa tracer for sap movement, but they are fundamentally differentin their operating principles. Two methods commonly employed,the stem heat balance and trunk sector heat balance methods,use the heat balance principle; the stem is heated electricallyand the heat balance is solved for the amount of heat takenup by the moving sap stream, which is then used to calculatethe mass flow of sap in the stem. In the heat-pulse method,rather than using continuous heating, short pulses of heat areapplied and the mass flow of sap is determined from the velocityof the heat pulses moving along the stem. In addition, ratesof sap flow can be determined empirically, using the thermaldissipation technique, from the temperature of sapwood neara continuously-powered heater implanted in the stem. Users mustunderstand the theory underlying each of these methods, so thatthey can select the method most appropriate to their applicationand take precautions against potential sources of error. Whenattempting to estimate transpiration by stands of vegetationfrom measurements of sap flow in individual plants, users mustalso select an appropriate sampling strategy and scaling method. Key words: Sap flow, transpiration, stem heat balance, heat pulse velocity, review     

Improvement of the heat pulse method for determining sap flow in trees

Abstract. The heat pulse method for determining sap flux in large woody sterns was modified for easier field operation. It uses the measurement of the time elapsed between heat pulse release by a line heater radially inserted in the stem, and the occurrence of maximum temperature 15 mm downstream of the heater. This spacing between heater and thermometer is critical to the reliability of the measurement. Calculations using uncorrected theory provide estimates of the sap flux density in stems with both uniform and non-uniform cross-sectional distribution of conducting tissues which are about 55% of the actual sap flux density. This factor results from insufficient thermal homogeneity between tissues where sap flow occurs and tissues where sap flow has been interrupted.
Sap flow in trunks of citrus trees was inferred from measurements of the cross-sectional distribution of sap flux density. Variability of sap flux density is specific to each trunk and is time-dependent and imposes multiple radial and angular measurements. The method was checked in a citrus trunk ramified into three branches. Instantaneous determinations of the flow in the trunk and in the branches differed by less than 5.7%. The daily values agreed within 2.8%.     

Changes in stem water content influence sap flux density measurements with thermal dissipation probes

Key message

Stem WC may decline during the day. Zero-flow dT _m increases when WC decreases. Use of nighttime dT _m in the calculation of sap flux density during the day might introduce errors.

Abstract

There is increasing evidence of diel variation in water content of stems of living trees as a result of changes in internal water reserves. The interplay between dynamic water storage and sap flow is of current interest, but the accuracy of measurement of both variables has come into question. Fluctuations in stem water content may induce inaccuracy in thermal-based measurements of sap flux density because wood thermal properties are dependent on water content. The most widely used thermal method for measuring sap flux density is the thermal dissipation probe (TDP) with continuous heating, which measures the influence of moving sap on the temperature difference between a heated needle and a reference needle vertically separated in the flow stream. The objective of our study was to investigate how diel fluctuations in water content could influence TDP measurements of sap flux density. We analysed the influence of water content on the zero-flow maximum temperature difference, dT _m, which is used as the reference for calculating sap flux density, and present results of a dehydration experiment on cut branch segments of American sycamore (Platanus occidentalis L.). We demonstrate both theoretically and experimentally that dT _m increases when stem water content declines. Because dT _m is measured at night when water content is high, this phenomenon could result in underestimations of sap flux density during the day when water content is lower. We conclude that diel dynamics in water content should be considered when TDP is used to measure sap flow.     

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     

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements

Summary Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day^–1 tree^–1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m² of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s^–1 that is equivalent to 112 mmol m^–2 s^–1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m^–2 s^–1 at the top and 150 mmol m^–2 s^–1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was –27.76±0.27 (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.This paper is dedicated to Prof. Dr. O.L. Lange on the occasion of his 65th birthday     

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.     

The accuracy of the thermal dissipation technique for estimating sap flow is affected by the radial distribution of conduit diameter and density

There are conflicting reports on the accuracy of the thermal dissipation probe (TDP, the Granier method) measurement using the original formula, which is widely used to estimate the transpiration of individual trees and forest stands. In this article, six woody species of three wood types were used to study a possible association between TDP measurement accuracy and wood anatomical characteristics, including the vessel diameter and density, as well as sapwood depth. We found that TDP technique with Granier’s original equation underestimated the sap flux density in six species to various degrees, dependent on conduit size and sap flux. Our calibration using two conifers with small diameters and a high density of tracheids was relatively consistent with Granier’s calibration; however, because there were larger diameters and lower densities of vessels in the two diffuse-porous species, the original calibration significantly underestimated sap flow. Two ring-porous species had the largest diameters and lowest densities of vessels. In particular, Robinia pseudoacacia possessed the shallowest sap wood depth, less than a probe length. Our calibration for the ring-porous species, especially R. pseudoacacia, deviated far from the original calibration, which mostly underestimated the sap flow. The degree of underestimation was well associated with sap wood depth and the radial diameter and density distribution of conduits. Our results demonstrated that a new calibration must be operated for each species together with the sapwood depth determination and more probes may be applied for one stem in the field to obtain the more accurate sap flux. In addition, we investigated the effects of different environmental temperature and perfusing fluid composition on the TDP-based sap flux measurement. We found that an environmental temperature reduction from 25 to 0 °C did not alter the values of the maximum temperature difference (ΔT_m) between a heated probe and a reference probe when there was no sap flow, verifying that ΔT_m measured at night can be used as a reference in daytime.     

A comparison of sap flux and water relations of leaves of various isolated trees with special reference to foundation movement in clay soil

Diurnal variation in sap flux (S) through stems of six trees, two each of Ulmus procera SALISB., Melaleuca styphelioides SM. and Prunus cerasifera EHRH. ‘Nigra’ (referred to hereafter by their generic names), were estimated from measurements of heat pulse velocities. Leaf water potential (ψ), stomatal conductance (g _s ) and transpiration from leaves (T) of all replicate trees were measured at 1300–1500h, once during the summer. On two separate occasions measurements were made of S, ψ, (g _s ) and T for one each of Ulmus and Melaleuca trees to study diurnal variations in these parameters. A 12×12 m² area around each tree was kept covered to simulate the condition of trees growing on pavements adjacent to residential properties. Sap flux for these tree species was in the order Melaleuca>Ulmus>Prunus. It is suggested that the smaller canopy and sapwood area in Prunus compared to the other two species is responsible for lower water potential and lower transpiration rate than the other species. Detailed analysis of the diurnal variation in sap flux and water relation of leaves of Melaleuca and Ulmus indicated sap flux of Melaleuca to be greater than that of Ulmus at the same transpiration rate per unit leaf area although the sapwood area of the two species was marginally different. This may have been due either to the difference in canopy conductance or in leaf area between the two species. With the assumption that sap flux closely resembles the rate of soil water extraction for both species, results indicate that Melaleuca is likely to extract soil water at a higher rate than Ulmus and hence is capable of causing greater shrinkage and soil movement than Ulmus.     

Validating sap flow measurement in field-grown sunflower and corn

Sap flow measurement has been widely used recently in studyingplant response to the environment, but results have not alwaysbeen satisfactory. The possibility that the non-uniform distributionof the conducting elements in the stem cross-sectional areaintroduces error in the measurements was examined. The heatpulse method, combined with anatomical observations of the stemat the point of sap flow measurement, was applied in corn andsunflower, to study the interaction between anatomical structureand sap flow distribution. The probability of the temperaturesensor being in contact with conducting elements was stronglyaffected by its insertion depth into the stem; the measuredheat pulse velocity for a given transpirational flux was correlatedwith the number of conducting elements in contact with the sensor.Consequently, the calibration coefficient is affected by theposition of the thermocouple junction in relation to the conductingelements. This dependence of apparent heat pulse velocity ondensity of conducting elements suggests that thermal equilibrationin species with large stems may not be achieved within the limitedtime of heat dissipation. The relationship between sap flowand leaf area of single plants was used to test the consistencyand the accuracy of the sap flow measurements. The ratio ofmeasured to potential transpiration was used to validate theuse under field conditions, of the calibration coefficient determinedin potted plants. Key words: Heat pulse, transpiration, LAI, potential transpiration     

Comparison of sap flow, cavitation and water status of Quercus petraea and Quercus cerris trees with special reference to computer tomography

Concurrent measurements of sap velocity (heat pulse) and ultrasound acoustic emission were performed on the trunks of mature Turkey oak (Quercus cerris) and sessile oak (Quercus petraea) trees. Plant water status was assessed by measuring leaf water potential, leaf conductance and transpiration. Wood density was estimated non-destructively on the trunk section of the plants by mobile computer tomography, which measures the attenuation of a collimated beam of radiation traversing the trunk in several directions, as the device rotates around the tree. Absorption is proportional to the density of the wood. As wood density is strictly correlated to water content, this non-invasive method allows the water content in the trunk section to be evaluated as well as mapped. Leaf water potential declined each morning until a minimum was reached at midday and recovered in the afternoon, lagging behind changes in transpiration rate. Good correspondence was found between the patterns of sap velocity and cavitation rate. A close correlation was demonstrated between wood density, water content and sap velocity. Sap now was always higher in Turkey oak than in sessile oak. Trunk signatures by computer tomography appeared to differentiate the two oak species, with the Turkey oak stem clearly more hydrated than the sessile oak; water storage reservoirs could play an important role in tree survival during extended periods of low soil water availability and in the relative distribution of tree species, especially in the context of global climate change. Late-wood conducting elements of oak species seem to play a significant role in water transport. The mobile computer tomograph was confirmed as a peerless tool for investigating stem water relations. Diurnal variations in the measured parameters under natural drought conditions and the differences between the two oak species are discussed.     

Sap flux density measurements based on the heat field deformation method

Accurate measurements of whole tree water use are needed in many scientific disciplines such as hydrology, ecophysiology, ecology, forestry, agronomy and climatology. Several techniques based on heat dissipation have been developed for this purpose. One of the latest developed techniques is the heat field deformation (HFD) method, which relies on continuous heating and the combination of a symmetrical and an asymmetrical temperature measurement. However, thus far the development of this method has not been fully described in the scientific literature. An understanding of its underlying principles is nevertheless essential to fully exploit the potential of this method as well as to better understand the results. This paper therefore structures the existing, but dispersed, data on the HFD method and explains its evolution from an initial ratio of temperature differences proportional to vapor pressure deficit to a fully operational and practically applicable sap flux density measurement system. It stresses the importance of HFD as a method that is capable of measuring low, high and reverse flows without necessitating zero flow conditions and on several sapwood depths to establish a radial profile. The combination of these features has not been included yet in other heat-based sap flow measurement systems, making the HFD method unique of its kind.     

荒漠防护林典型树种液流特征及其对环境因子的响应

利用基于热补偿理论的SF300分体液流仪对干旱荒漠区人工防护林典型树种(俄罗斯杨、胡杨、榆树、沙枣)树干液流全天候监测,自动气象站同步记录相关环境因子变化。研究表明:①4种防护林树种茎干液流日变化除沙枣树外均存在明显昼夜节律,液流速度在同属种间差异较小,在不同属种间差异显著,俄罗斯杨的日平均液流速度可以达到沙枣的13.8倍,耗水量排序为俄罗斯杨胡杨榆树沙枣树。②水分充足条件下,增加实验地灌溉量使4种树木蒸腾受到抑制,液流流速降低,水分利用效率降低。③液流流速因所处树干径向位点不同而存在差异,俄罗斯杨、榆树、沙枣液流速度表现出由形成层到髓心的递减趋势,胡杨树干径向位点液流没有表现一定规律。④树干液流流速与环境因子进行相关分析,通过逐步回归分析建立了4个典型树种茎干液流速度与环境因子关系估算模型,分析认为4种树木的环境敏感性排序为俄罗斯杨榆树胡杨沙枣。     

The effect of fertilization on sap flux and canopy conductance in a Eucalyptus saligna experimental forest

Land devoted to plantation forestry (50 million ha) has been increasing worldwide and the genus Eucalyptus is a popular plantation species (14 million ha) for its rapid growth and ability to grow well on a wide range of sites. Fertilization is a common silvicultural tool to improve tree growth with potential effects on stand water use, but the relationship between wood growth and water use in response to fertilization remains poorly quantified. Our objectives in this study were to determine the extent, timing and longevity of fertilization effects on water use and wood growth in a non‐water limited Eucalyptus saligna experimental forest near Hilo, HI. We evaluated the short‐ and long‐term effects of fertilization on water use by measuring sap flux per unit sapwood area, canopy conductance, transpiration per unit leaf area and water‐use efficiency in control and fertilized stands. Short‐term effects were assessed by comparing sap flux before and after fertilizer application. Long‐term effects were assessed by comparing control plots and plots that had received nutrient additions for 5 years. For the short‐term response, total water use in fertilized plots increased from 265 to 487 mm yr^?1 during the 5 months following fertilization. The increase was driven by an increase in stand leaf area accompanied by an increase in sap flux per unit sapwood area. Sap flux per unit leaf area and canopy conductance did not differ during the 5 months following fertilizer additions. For the last 2 months of our short‐term measurements, fertilized trees used less water per unit carbon gain (361 compared with 751 kg H₂O kg C^?1 in control stands). Trees with 5 years of fertilization also used significantly more water than controls (401 vs. 302 mm yr^?1) because of greater leaf area in the fertilized stands. Sap flux per unit sapwood area, sap flux per unit leaf area, and canopy conductance did not differ between control and fertilized trees in the long‐term plots. In contrast to the short‐term response, the long‐term response of water use per unit wood growth was not significant. Overall, fertilization of E. saligna at our site increased stand water use by increasing leaf area. Fertilized trees grew more wood and used more water, but fertilization did not change wood growth per unit water use.     

黄土丘陵区辽东栎和刺槐树干液流时滞效应与蒸腾特征的关联性

运用Granier热扩散探针法,于2016年7-9月对半干旱黄土丘陵区天然次生林树种辽东栎和人工林树种刺槐的树干液流进行连续测定,并同步监测气象因子和土壤含水量,用错位相关法分析液流通量密度与空气水汽压亏缺日变化的时滞长度,研究2个树种不同径级个体在不同土壤水分条件下液流通量密度与蒸腾驱动因子之间的时滞效应.结果表明:辽东栎和刺槐液流通量密度的日变化节律与气象因子显著相关,空气水汽压亏缺峰值的出现较辽东栎树干液流通量密度滞后118.2 min,较刺槐树干液流通量密度滞后39.5 min;而光合有效辐射的峰值通常滞后于辽东栎12.4 min,提前于刺槐68.5 min.液流通量密度和空气水汽压亏缺的时滞长度与树种和土壤含水量显著相关,辽东栎、刺槐在土壤含水量较高时段的时滞长度分别大于土壤含水量较低时段32.2和68.2 min.时滞长度与径级的相关性整体上未达到显著水平,但在土壤含水量较低时段小径级刺槐的时滞长度大于大径级21.4 min,差异达到了显著水平.两树种液流通量密度与空气水汽压亏缺之间的时滞效应反映了对蒸腾驱动因子的敏感性,较好的土壤水分条件有利于液流通量密度提早达到峰值,较低土壤水分会导致树干液流对气象环境因子响应的敏感性降低;刺槐树干液流受土壤水分的影响更显著.     

热脉冲技术3种方法组合在测量树干液流中的应用

利用树干液流方法获取树木蒸腾特征对理解树木水分生理、森林生态和森林系统水分交换具有重要意义.利用广泛应用于土壤热参数和土壤蒸发测量的
三针热脉冲探头,基于热比率法(HRM)、最大温度法(T_Max)和单针热脉冲法(SHPP)同时实现了旱柳液流密度的测定,并与热扩散探针(TDP)测量结果进行对比分析.结果表明： 三针热脉冲探头安装约5周后进入稳定测量阶段,3种方法初期测量结果比稳定测量阶段高135%~220%,HRM、T_Max和SHPP法与TDP测量结果具有显著的线性相关性,R2分别为0.93、0.73和0.91,SHPP与HRM法测定结果的R²达到0.94.HRM在低速和逆向液流时测量具有较高的精度;SHPP探头配置简单、测量精度高,但无法甄别液流方向,是测定液流非常有前途的方法;T_Max测量液流误差较大,无法测量<5 cm³·cm^-2·h^-1的液流,不建议单独用于液流测量,但其能够准确测定树干热扩散系数,并可用于其他方法液流计算.建议根据试验目的,选取不同方法或者几种方法组合进行树干液流测量.     

Evaluation of sap flow methods to determine water use by cultivated palms

Sustainable management of water resources allocated to palm crops requires reliable tools for measuring palm water use. Thermometric sap flow methods developed for woody dicot plants hold great potential for use in palms, but there have been few investigations to determine whether such methods can be used successfully in ‘woody’ monocots. Here, we evaluate two sap flow methods for measuring whole-plant water use by potted cocos palms (Syagrus romanzoffiana): the Heat Ratio Method (HRM) and the Compensation Heat Pulse Method (CHPM). Measurements of whole-plant water use from the HRM and the CHPM were compared to gravimetric measurements acquired from an electronic balance. Of the two methods, the HRM gave the most accurate results and it most precisely described patterns of transpiration with respect to environmental conditions and leaf morphology. Estimates of total daily water use from the HRM and the balance were highly correlated (R² = 0.92; P < 0.0001) and very near to a 1:1 relationship—an excellent result given the potential for error associated with each method. As expected, the CHPM was seriously limited at low flow rates, but it agreed well with the HRM at higher flow rates (heat pulse velocity >4 cm h⁻¹). Anatomical investigations revealed that vascular bundles in measured palm fronds were evenly distributed and the distance between bundles was comparable in scale to the diameter of probes of HRM sensors, and most likely an order of magnitude smaller than the probable zone of thermal influence for HRM measurements. This contention was supported by results of mathematical modeling suggesting that HRM heat pulse velocities in palm-like sapwood remain largely unaffected by increases in heterogeneity caused by larger xylem vessels and wider interstitial tissues. Although wounding models for heat pulse sap flow methods have not historically catered for the effects of discrete vascular bundles, they are sufficiently approximate given our empirical validations, and we conclude that palm sapwood is thermally homogenous enough for the HRM and the CHPM to be used without modification. Overall, we present strong evidence that heat pulse methods such as the HRM and the CHPM can be used successfully in woody monocots. Finally, we note that the CHPM is likely to perform better in cases of very high flow rates, while the HRM provides more comprehensive monitoring of the usual range of flow rates including at night, early morning and during water deficit.     

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.