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%.     

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.     

科尔沁沙地黄柳和小叶锦鸡儿茎流及蒸腾特征

利用Dynamax茎流测量系统、Li-6400光合作用仪和自动气象站分别对科尔沁沙地两种灌木树种黄柳和小叶锦鸡儿的液流变化、蒸腾速率及其周围的环境因子进行检测.研究结果表明：（1）黄柳、小叶锦鸡儿茎干液流通量密度日变化趋势基本相同,呈多峰曲线;液流启动时间分别为4：30、5：30,在13：00左右到达各自液流峰值,峰值大小为81.2～91.7mg/h、17.3～27.1mg/h,20：30降为最低,晚间均具有明显的液流活动现象.（2）灌木的茎流日变化曲线与蒸腾速率日变化曲线并不吻合,实验测得的叶片蒸腾速率不能同步反映茎干液流的动态变化特征;黄柳的叶片蒸腾速率和单枝蒸腾耗水量均大于小叶锦鸡儿,耐旱性相对较低.（3）黄柳、小叶锦鸡儿白天液流通量密度变化趋势与环境因子变化趋势相吻合.相关性分析表明,影响灌木液流变化的主要因子依次是太阳有效辐射、相对湿度、大气温度、风速、10cm和20cm土壤温度.     

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.     

An external heat pulse method for measurement of sap flow through fruit pedicels, leaf petioles and other small-diameter stems

The external heat ratio method is described for measurement of low rates of sap flow in both directions through stems and other plant organs, including fruit pedicels, with diameters up to 5 mm and flows less than 2 g h⁻¹. Calibration was empirical, with heat pulse velocity ( v _h) compared to gravimetric measurements of sap flow. In the four stem types tested ( Actinidia sp. fruit pedicels, Schefflera arboricola petioles, Pittosporum crassifolium stems and Fagus sylvatica stems), v _h was linearly correlated with sap velocity ( v _s) up to a v _s of approximately 0.007 cm s⁻¹, equivalent to a flow of 1.8 g h⁻¹ through a 3-mm-diameter stem. Minimum detectable v _s was approximately 0.0001 cm s⁻¹, equivalent to 0.025 g h⁻¹ through a 3-mm-diameter stem. Sensitivity increased with bark removal. Girdling had no effect on short-term measurements of in vivo sap flow, suggesting that phloem flows were too low to be separated from xylem flows. Fluctuating ambient temperatures increased variability in outdoor sap flow measurements. However, a consistent diurnal time-course of fruit pedicel sap flow was obtained, with flows towards 75-day-old kiwifruit lagging behind evaporative demand and peaking at 0.3 g h⁻¹ in the late afternoon.     

应用热平衡法测定玉米/大豆间作群体内作物的蒸腾量

通过田间试验采用基于热平衡法的茎流计测定玉米/大豆条带间作群体内作物的蒸腾规律.结果表明：间作群体内,玉米和大豆植株的茎流速率在晴天呈单峰曲线,在阴天则呈多峰曲线.植株的茎流受多个环境因子的影响,其中太阳辐射是影响植株茎流最主要的气象因子.玉米和大豆的单株日茎流量与多个气象因子间存在较好的相关关系,达到极显著水平.茎流观测期内（2008年6月1-30日）,间作群体内玉米植株的日均蒸腾量（1.44 mm·d^-1）为大豆（0.79 mm·d^-1）的1.8倍,玉米和大豆植株的蒸腾量分别占间作群体总蒸腾量的64%和36%.考虑到作物的茎直径和叶面积的空间变异,安装一定数量的茎流探头对于准确测定植株茎流是十分必要的.     

Sap Flow of Populus euphratica in a Desert Riparian Forest in an Extreme Arid Region During the Growing Season

In the present study, the heat pulse technique was applied to investigate the stem sap flow of Populus euphratica in a desert riparian forest in an extreme arid region from April to October 2003 and from May to October 2004. The experimental sites were in Qidaoqiao （101 °10′ E, 41°59′ N） and Bayantaolai farm （101°14′ E, 42°01′ N） in Ejina county, in the low reaches of the Heihe River, China. The results indicated that the diurnal change in the velocity of sap flow showed minor fluctuations. At night, the rising of sap flow could be observed in the main tree species because of root pressure. During the growing season, the maximum average velocity was observed in July, followed by August, and the same velocity was observed in September and May; the minimum velocity was observed in October. The transpiration from June to August during the growing season accounted for approximately 70% of the annual total transpiration. The sap flow velocity of P. euphratica trees of different ages could be arranged in the order： 15 yr 〉 25 yr 〉 50 yr. Sap flow velocity was closely related to changes in micrometeorological factors, with average sap flow velocity showing a significant linear correlation with net radiation, air temperature and relative humidity.     

An Improved Thermoelectric Probe for Measurements of Apparent Sap Flow Velocity in Intact Plant Stems

Gray, H. R-, Erickson, P. I. and Stone, J. F. 1985. An improvedthermoelectric probe for measurement of apparent sap flow velocityin intact plant stems.—J. exp. Bot. 36: 1320–1324. An improved thermoelectric technique for non-intrusive, non-destructive,and direct sensing of the apparent velocity of sap flow in intactplant stems is presented. The thermoelectric probe was madeof extruded Teflon (DuPont) rod. One of two heating elements,positioned 0.5 cm and 1.0 cm below the temperature sensor, wasused to generate the heat pulse. Heating element selection wasbased on daily environmental conditions. A thermistor touchingthe stem sensed the heat pulse. A second thermistor mountedin dose proximity was used to cancel ambient temperature fluctuations. The improved thermoelectric probe was tested using irrigatedpeanuts (Arachis hypogeae L.). After canopy closure was completein both treatments, apparent sap flow velocity(V_a) was measuredon 17 dat 1200 s intervals between 0830 h and 1630 h apparentsolar time. Successful V_a, measurements were accomplished on94% of the attempts. In order to verify performance on otherspecies with different types of stems, non-replicated V_a, determinationswere also made on cotton (Gossypiwn spp.), prickly sida (Sidasptnosa L.), goosegrass (Eleusine indica (L.) Gaertn.), Palmerpigweed (Amaranthus Palmerii S. Wats.), and cottonwood (Populusdeltoides Marsh.). Success of the probe was due in part to the design of the probewhich included remotely selectable spacing between heater andthermistor and adequate insulation and shielding of the probeand sensed portion of the stem. In addition, acclimatizationof the probe to the operating environment was essential. Key words: Sap velocity, instrumentation     

EFFECT OF SOIL MOISTURE STRESS ON THE CORRELATION BETWEEN HEAT PULSE VELOCITY AND TRANSPIRATION

(1) Heat pulse velocity in the stem of pine and sour orangeseedling, measured by HUBER's heat pulse method, was correlatedwith transpiration as measured by determining the humidity ofair passed over the plant. This correlation was used to calculatetranspiration under natural conditions, when heat pulse velocityonly was measured. (2) It was shown that heat pulse velocity at a given transpirationrate was related to water stress. When water potential in thesoil decreased heat pulse velocity increased in pine seedlingsand decreased in sour orange. This effect of water stress didnot appear to be completely reversible. Under conditions ofvery low water potential in the soil, no useful correlationbetween transpiration and heat pulse velocity could be obtainedeither in pine or sour orange. (3) From the data obtained it is concluded that transpirationcan be reliably calculated from heat pulse velocity measurementsonly if the calibration curve relating transpiration to heatpulse velocity is determined shortly beforehand under similarconditions of soil moisture and potential evapotranspiration.Even so, only total daily transpiration can be estimated bythis method. Hourly figures of transpiration are unreliable,especially under conditions conducive to low water potentialin the plant. (Received June 24, 1964; )     

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.     

Laurel forests in Tenerife, Canary Islands

The efficiency of the conductive system in about 40-year-old Laurus azorica trees growing in a laurel forest was evaluated by comparing main stems and leaves (petioles) on the basis of theoretical sap flow values (1) calculated from vessel anatomy (taking vessels as ideal capillaries), (2) derived from measured dye velocity and (3) data taken from direct sap flow measurements. It was found that actual sap flow rate per wood area increases in stems from the pith towards the cambium. The outermost part of the stem is the most important part of the tree for conducting water. Maximum actually measured transpiration (sap flow rate) for the stand was practically identical to the theoretical rate calculated based on petiole anatomy, but it was about 45 times lower than that calculated based on stem anatomy. This illustrates the safety features of stem wood, which due to its high vessel density, is capable of transporting all the water required even when only a small area of its vessels is working. In the petioles, xylem is more efficiently used, but almost all vessels must work in order to supply water to leaves and any disturbance may cause leaf loss.     

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.     

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。     

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.     

小叶锦鸡儿灌丛群落蒸腾耗水量估算方法

为了确定沙地小叶锦鸡儿 (Caragana microphylla) 人工林的蒸腾速率, 于2006年6月运用热平衡茎流测量技术, 对科尔沁沙地一处15a生人工小叶锦鸡儿群落的分枝液流动态进行了监测。根据生物统计结果选取被测标准枝, 标准枝基径在0.4~1cm范围内。同时, 分别用基径总断面积推算法和叶片密度推算法对灌丛叶面积进行了估算。以叶面积为扩展纯量, 利用标准枝液流对灌丛群落耗水量进行尺度转换, 在转换过程中, 假设叶面积与蒸腾耗水量之间具有很强的相关性。该尺度转换方法经与大型称重式Lysimeter测值对比验证, 误差小于14.3%, 可望准确估算小叶锦鸡儿灌丛群落的蒸腾耗水量。     

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     

植物夜间液流的发生、生理意义及影响因素研究进展

植物夜间液流是指在夜间通过植物根、茎、叶的液流量。通过对不同物种、生境条件和生态系统的野外观测,发现植物普遍存在夜间液流现象。阐述了夜间液流的大小和组成,并从夜间液流的生理意义、影响因素以及生态水文效应方面对已有的研究进展进行了综述和分析。夜间液流占到全天液流量的比例一般为5%—20%。夜间液流包括夜间的茎干补水和夜间的蒸腾作用两个过程,但是目前没有确切的研究或技术将两个过程区分开来。虽然总体上夜间液流占全天液流量的比例较少,但是夜间液流的储水作用和蒸腾作用对植物生长有重要的生理意义:夜间储水作用提高了夜间茎干水势,减少了木质部栓塞化的形成,加强了植物对干旱环境的适应;而蒸腾作用在营养物质和氧气的运输,以及水力提升等方面有重要的作用。影响夜间液流的因素较多,气象因素是主要的环境驱动因子,而土壤水分对夜间液流的影响与生境有关;夜间液流还受到物种和生境条件的影响。由于夜间液流的发生,对不同尺度的生态水文过程产生了影响。未来的研究可进一步探索在全球气候变化条件下,夜间液流与植物生理过程的关系,定量评估夜间液流对生态水文过程的影响,深入研究夜间液流对环境变化的响应。     

Heat balance sap flow gauge for small diameter stems

Applying heat balance sap flow gauges to plant stems <10 mm in diameter has been difficult because a miniature design is needed that can be attached to a range of stem geometries. This report presents a modified gauge design for use on small plant stems of irregular geometry and shows results from Glycine max with stem diameters of 3–4 mm. The gauge was evaluated on container-grown plants by comparing gauge measurements of flow to gravimetric estimates of transpiration. Experiments were conducted in the laboratory and greenhouse, using artificial and natural lighting, respectively. Laboratory comparisons of gauge versus gravimetric water loss measurements indicated that the instrument was accurate to within ±5% when soil water was not limiting. Similar results were obtained from greenhouse tests except when soil water availability was low and cumulative gauge estimates became 30–45% less than gravimetric measurements. Differences may have reflected reduced plant water uptake or errors in sap flow estimates associated with low flow rates. Gauge accuracy was not improved by including the rate change in heat storage (S) in the flow calculations because S was always less then 3% of the total heat balance. Relationships between system temperature and sap flow rate suggested a diagnostic test for determining optimum power input. A time constant of 15 s indicated potential application in many agronomic and physiological studies.