Sap flow measurements with some thermodynamic methods,flow integration within trees and scaling up from sample trees to entire forest stands

Sap flow measurement techniques and evaluation of data are reviewed. Particular attention is paid to the trunk segment heat balance (THB) and heat field deformation (HFD) methods based on 30 years experience. Further elaboration of sap flow data is discussed in terms of integrating flow for whole stems from individual measuring points, considering variation of radial patterns in sapwood and variation around stems. Scaling up of data from sets of sample trees to entire forest stands based on widely available biometric data (partially on remote sensing images) is described and evaluated with a discussion of the magnitude of errors, the routine procedure applicable in any forest stand and practical examples.     

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%.     

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.     

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     

树干自然温度梯度变化对热扩散法测算树干液流速率的影响

为了揭示树干自然温度梯度的变化规律及其对树干液流速率测算结果的影响,于2007年5月至10月利用改进的SF-L型热扩散式树液流测定装置,对北京低山区生长的油松和侧柏的树干自然温度梯度、加热温差和气象、土壤水分因子进行了连日同步监测。结果表明:(1)树干自然温度梯度对加热针温差的影响,侧柏大于油松。(2)树干自然温度梯度对液流速率计算结果的影响具有显著的统计意义(P0.01),平均误差大于30%,误差峰值出现在太阳高度角较小的时候。(3)影响树干自然温度梯度最重要的环境因子是光照强度,其次是空气温度。上述结果说明,树干自然温度梯度对热扩散法测定的液流速率的影响不可忽视,研究树木耗水机制时应予以充分考虑。光照强度和空气温度是影响树干自然温度梯度最重要的两个因子,但其影响机制仍需进一步研究。     

Sap Flow Measurements in a Socotra Dragon’s Blood Tree (<Emphasis Type="Italic">Dracaena cinnabari</Emphasis>) in its Area of Origin

For the first time, in situ field measurements of sap flow were conducted in adult Dracaena cinnabari plant native to the arid tropical climate of Socotra Island. The heat field deformation (HFD) method was applied using both single and multi-point sensors to study azimuthal and radial sap flow variability in stem, roots and first-order branches over two weeks during a winter monsoon. The main aim of this work was to monitor sap flow in adult D. cinnabari in-situ to better understand its physiological adaptation to extreme arid environments. The second aim was to compare our results with earlier sap flow measurements in adult D. draco uing the same HFD method. The last question we wanted to answer was comparison of sap flow measurements in both, young and adult Dracaena species. We found that sap flow magnitude is low and of a similar range in all observed D. species. High sap flow variability was recorded in different parts of adult D. cinnabari plant which changed throughout the day responding to interplay between intrinsic and extrinsic water potential gradients induced by sunlight. Maximum sap flow levels had variable pattern around stem in response to sun exposure, similarly as it was observed in adult D. draco plant. Sensors installed tangentially in stem xylem showed that water transport in adult D. cinnabari may move in lateral direction. This work also presents several methodological aspects detected from earlier observations of dicots which proved to be more pronounced in adult D. species. These methodologies relate to interpreting negative sap flow rates in conjunction with established axial flow reversal during hydraulic redistribution usually occurring under low evaporative demands and dry soil. Conversely, flow reversal during the day under high evaporative demands and wet soil may designate lateral water movement induced by internal water redistribution.     

五针热脉冲探头在测定树干液流中的应用

准确测量树干液流对研究树木耗水特性、植物生理和生态水文效应等具有重要意义.本文选用一种新型五针热脉冲多功能数字探头(PHPP),运用热脉冲理论,测量小叶杨树干液流,并与热扩散探针(TDP)进行对比分析,探究五针热脉冲探头测量树干液流的适用性和准确性.结果表明: 五针热脉冲探头能够准确揭示小叶杨树干液流昼夜变化规律,并且与热扩散探针测量结果呈显著的线性相关关系,R²达到0.90,均方根误差为2.75,平均相对误差为11%.PHPP探头能较为精确地识别低液流和逆液流,可以快速准确地拟合热参数,直接测得树干液流速率,应用潜力较大.     

Sap flow measurements in grapevines (Vitis vinifera L.) 1. Stem morphology and use of the heat balance method

The heat balance method was evaluated in detail for its use in older, mature grapevines with stems of 35 – 45 mm in diameter. Dye colouring of the xylem vessels revealed that even 21 year old grapevines did not show any development of heartwood and that xylem vessels of that age still have the capacity to transport water. A comparison of weight loss of potted vines on a balance and sap flow measurements demonstrated that the heat balance system reflected rapid changes in flow rate without any time delay. However, since even 20 year old xylem vessels of grapevines have the capacity to conduct water, the heater band was not able to heat the sap in all year rings evenly. Apparently, at low flow rates this effect was small and sap flow was calculated correctly. With increasing flow rates large thermal heterogeneities developed upsetting the calculation of the heat balance and mass flow. Consequently, actual sap flow was overestimated by 50 to 100% at high flow rates. This could be attributed to thermal gradients in these relatively thick stems excluding the use of this technique for measurements of long term as well as short term water use patterns in older grapevines. This revised version was published online in June 2006 with corrections to the Cover Date.     

Instrumental methods for studies of structure and function of root systems of large trees

New methods using different physical principles have been successfully applied in studies of root systems of large trees. The ground-penetrating radar technique provides 3D images of coarse roots (starting with a diameter of about 20 mm) from the soil surface down to a depth of several metres. This can even be done under layers of undisturbed materials such as concrete, asphalt and water. Fine roots cannot be visualized by this method, but the total rooted volume of soil can be determined. The differential electric conductance method has been used for fast measurement of conducting (absorbing) root surfaces. However, more testing is needed. Both these methods are non-invasive. The results can be verified by an almost harmless excavation of whole root systems, including fine roots, using the ultrasonic air-stream (air-spade) method. This method is suitable for all studies, as well as practical operations on roots or objects in their vicinity, where a gentle approach is required. Sap flow measurements on their own or in tandem with soil moisture monitoring play a leading role in studying root function and hydraulic redistribution of flow in the soil. The water absorption function of roots can be studied by measuring sap flow on individual root branches directly (as on crown branches) and also indirectly, by measuring the radial pattern of sap flow in different sapwood depths at the base of a stem. Root zone architecture can also be estimated indirectly by studying its functionality. The heat field deformation method with multi-point sensors has been found to be very convenient for this purpose. A combination of several such methods is recommended whenever possible, in order to obtain detailed information about the root systems of trees.     

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     

On-line Estimation of the Rate of Sap Flow in Plant Stems Using Stationary Thermal Response Data

The heat pulse method for the measurement of the rate of sapflow in plants suffers from two drawbacks, viz. the discontinuousnature of the measurements and the lack of a precise mathematicalformulation of the heat fluxes. To overcome these drawbacksa random noise heat signal has been used as an input and twosensor pairs have been used for the measurement. A model isgiven that employs the signal from one sensor pair as inputand the signal from the second pair as output of a heat transportsystem. Thus the adverse effects of radial heat conduction havebeen eliminated. Simulation models with the rate of sap flowas one of the parameters have been fitted to the stationaryresponse data with an on-line parameter optimization algorithm.The method has been validated in applications to artificialdata and to real data obtained from a plastic tube and fromtomato plants. Key words: Sap flow, Heat pulse method, Continuous measurement     

应用热技术研究树干液流进展

综述了热技术方法测定树干液流的基本原理和不同适用范围．通过各种热技术方法可以确定树干水分运输格局及其数量;与树木生理指标和环境因素联合测定,可深入分析整树水分导度、气孔导度、边界层导度、水势及树干储水与树木蒸腾之间的关系,探讨树干液流受外界环境因素影响程度及其响应,揭示树木蒸腾内在的调节机制和外在影响因素;热技术可用于长期连续测定地带性森林主要树种蒸腾耗水特征。为正确评价森林的水文效应提供技术支持．     

应用Granier热消散探针（TDP）法测定树木的木质部液流：理论与实践

Granier热消散探针法是目前研究树木生理生态和森林水文最常用的测定整树水分利用的方法之一。然而,已有的相关综述文献中,还没有专门介绍利用Granier热消散探针研究木质部液流的综述文章。本文重点介绍了Granier热消散探针测定系统的理论基础,对有关该探针的校准和改进的最新研究进展进行了综述,并还深入探讨了零液流信号值的确定、自然热梯度、损伤效应、液流的逆格型和将木质部液流密度外推至整树蒸腾耗水量等重要的实际问题。     

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.     

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     

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。     

不同天气条件下小叶锦鸡儿茎流及耗水特性

采用以热平衡为原理的Dynamax包裹式茎流测量系统,于2006年对科尔沁沙地人工小叶锦鸡儿群落的液流变化进行连续监测,利用实验区自动气象站同步检测光合有效辐射、气温、相对湿度和风速等气象因子,结合人工记录的天气状况,分别选取4种最典型的天气条件,分析了小叶锦鸡儿的茎流变化、单枝耗水量及其与气象因子的关系.结果表明:在高温、强辐射条件下,小叶锦鸡儿的液流呈"几"字型宽峰曲线,以低耗水的蒸腾特性耐受干旱缺水的环境.不同天气条件下,小叶锦鸡儿液流日变化波动曲线及其主要影响因子各异,但光合有效辐射始终是液流变化的主导因子.小叶锦鸡儿茎干液流通量密度的变化是各种气象因子综合作用的结果.单枝日耗水量的总体变化趋势是从晴天、阴天、风天到雨天依次减少,其平均值分别为459、310、281和193mg.d-1.     

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.     

干旱区枸杞树干液流变化特征及其影响因素

树干液流作为植物蒸散作用的水分来源,是植物水分消耗的直观量化监测指标,利用包裹式树干液流监测技术获取干旱区枸杞全生育期树干液流实时数据,分析了不同时间尺度树干液流变化特征及各气象要素对树干液流的影响,为明晰枸杞耗水规律及其影响因素提供了重要的佐证。结果表明:枸杞的树干液流量昼夜差异较大,白天液流量是夜间的10倍左右;晴天液流速率、日累积量及变化幅度均大于阴雨天气,晴天液流速率变化曲线且呈宽峰型,在06:30左右启动较阴天提前30min;夏季树干液流启动时间为6:00比秋季提前1h左右,夏季的峰值123g/h。盛果期液流速率最大10.32g/h,营养生长期最小1.35 g/h;6—8月旺盛生长季,平均日耗水1388.3g/d,5—11月全生育期日均耗水1102.7g/d;树干液流速率与太阳辐射、空气温度均呈极显著正相关关系,与相对湿度呈负相关关系;枸杞树干液流(F)与太阳辐射(S)、温度(T)、相对湿度(H)及饱和水汽压(VPD)符合方程F=41.5+0.167S-0.563H+1.36T-9.67VPD(R~2=0.6547)。     

A new method of sap flow rate determination in trees

A new method of sap flow rate determination in stems of adult trees is described in which mass flow (the transpiration flux in the present case) is estimated from a heat-transfer measurement. The device used includes a couple of thermometers in a differential connection and plate electrodes, through which a controlled heat input flows into a denned studied segment of the xylem. The present method was tested first with a laboratory simulating system, then with the stem of a living adult treePrunus avium L. The data are registered continually and automatically; continuous measurement of sap flow rate of long duration with a greater number of trees is thus made possible.