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%.     

黄土丘陵区辽东栎树干液流特征对边材面积和土壤水分的响应

运用Granier热扩散探针法对半干旱黄土丘陵区不同胸径辽东栎进行树干液流测定,并对太阳辐射、空气温湿度、降水量、土壤水分等环境因子进行同步观测,分析不同土壤水分条件下不同胸径辽东栎的树干液流变化特征及其对环境因子的响应.结果表明:辽东栎液流日变化特征总体上与太阳辐射和空气水汽压亏缺呈相同趋势,但液流峰值出现时间早于两个气象环境因子的峰值时间.同一树木个体在土壤水分条件较高时期的树干液流通量高于土壤水分较低时期.在相同土壤水分条件下,大径级样本液流通量显著高于小径级样本.采用指数饱和曲线函数对液流通量与太阳辐射和空气水汽压亏缺以及两因子的综合指标进行拟合,效果良好,可以反映液流通量对气象环境因子的响应规律.不同胸径辽东栎在不同土壤水分条件下的拟合曲线特征和拟合参数差异表明,在土壤水分较高时段,液流通量可快速上升至饱和值;在土壤水分较低时段,液流通量上升缓慢.小径级样本对土壤水分变化的反应更加剧烈.单位空气水汽压亏缺的日液流通量值(日液流通量与空气水汽压亏缺的比值)在两种土壤水分条件下的比值与边材面积呈线性相关,且小径级样本的斜率高于大径级样本,说明小径级样本对土壤水分的变化较为敏感,在土壤含水量较低时段,大径级样本较厚的导水组织对土壤水分供应不足起到了缓冲作用.     

The Effect of Hydrostatic Pressure Gradients on the Movement of Potassium across the Root Cortex

Transpiration reduces the hydrostatic pressure in the xylemand water moves into the root and across the root cortex inresponse to the pressure gradient so produced. This effect wassimulated by raising the pressure on the external medium surroundinga detopped root system. The flux of sap from the stem stumpand the concentration of potassium in the sap were measuredand the flux of potassium into xylem obtained as (sap flux Xconcentration of potassium). The application of a pressure of 2 atm. caused an approximatelyfourfold increase in potassium flux. This increase was independentof the presence of potassium in the external medium and wasdue, therefore, to an efflux of ions already stored in the roottissues. The effect was specifically due to hydrostatic pressureand not to the D.P.D. difference across the cortex, and wascaused, in part at least, by an increase in the permeabilityof the tissues to iona. This is evident since the increasedpotassium flux occurred in response to pressure, even when theconcentration of potassium in the xylem increased at the sametime, thus precluding any ‘dilution effects‘. It was confirmed that metabolic inhibitors and low temperaturereduce the sap flux but leave the concentration of the sap unchanged.Under a pressure gradient, however, there was an even greaterporportionate reduction of sap flux by equivalent concentrationsof inhibitor or low temperature and the concentration of potassiumin the sap gradually increased. Adding calcium or magnesium ions to the medium caused an increaseof potassium concentration in the sap, demonstrating exchangeprocesses in the transpiration pathway. The difficulty of framing a hypothesis to cover all the factsis discussed and it is tentatively suggested that movement acrossthe cortex to the xylem is a catenary process in which soluteand water move independently at one stage and together as asolution at another stage.     

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.     

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.     

热消散探针在粉单竹液流研究中的应用

通过对计算树干液流密度的Granier经验公式进行试验验证和校正,比较不同长度的热消散探针(TDP)测量粉单竹液流的结果,探讨TDP在粉单竹液流研究中的适用性,并与邻近生长的常见树种荷木的液流日变化进行比较分析.结果表明:由于粉单竹的竹壁较薄、解剖结构异质性较高,10 mm探针可能低估粉单竹的液流密度,8和5 mm探针监测的液流密度值较准确.粉单竹液流密度平均值和日变化格局的偏度均大于荷木,而夜间荷木的液流活动比粉单竹活跃,说明粉单竹的夜间补水不如荷木.以TDP探针研究竹子的液流是可行的,但应用于不同竹种之前,必须对Granier经验公式进行有效的验证和校正.     

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.     

甘肃石羊河流域干旱荒漠区花棒蒸腾耗水量

研究了甘肃石羊河流域干旱荒漠区天然生长条件下花棒的蒸腾耗水规律．结果表明：花棒木质部液流速率随探针插入深度的加大呈“高-低”变化趋势;主根直径较小的花棒各位点的平均液流速率上升速度较快,变幅较大;不同主根直径花棒的液流量相差较大,但变化趋势较为一致,即昼夜变幅较大,夜间液流量较小,白天液流量较大,呈多峰曲线;日液流量与蒸发蒸腾量ET0呈线性相关,蒸腾耗水主要在6-9月,占生长季总蒸腾量的79．04％;花棒生长后期日液流量与0—50cm深沙层含水量呈显著相关,与其它层含水量无明显相关性;气象因素对花棒树干液流量影响的大小表现为日均气温〉空气水汽压差〉风速．     

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.     

甘肃石羊河流域人工种植长穗柽柳树干液流量变化规律研究

采用SF200热脉冲液测定仪对人工种植长穗柽柳的树干液流进行连续监测;利用管式-时域反射仪测定长穗柽柳根区附近3m深土壤含水量,并用自动气象站对太阳辐射,空气温度,相对湿度,风速等主要气象因子进行同步监测,基于上述试验数据,分析长穗柽柳树干液流量的日季变化规律及与环境因子的关系。结果表明,人工种植环境下,霜冻对长穗柽柳树干液流量的影响较大,随着时间的推移,长穗柽柳日液流量可能会因环境因子的变化而上下波动,但总体上是逐渐减少的趋势;气象因子对长穗柽柳日液流量的影响依次为：太阳辐射〉气温〉空气水汽压差〉大气相对湿度〉风速。     

Xylem Sap Flux Affects Conventional Stem CO2 Efflux Measurements in Tropical Trees

The relationship between sap flux and stem CO₂ efflux was assessed for three mango trees. We observed higher than expected CO₂ effluxes at the place of measurement under intermediate sap flux velocities and lower fluxes under high sap flux velocity. This variation disappeared after removing the tree crown.     

Whole tree xylem sap flow responses to multiple environmental variables in a wet tropical forest

In order to quantify and characterize the variance in rainforest tree physiology, whole tree sap flow responses to local environmental conditions were investigated in 10 species of trees with diverse traits at La Selva Biological Station, Costa Rica. A simple model was developed to predict tree sap flow responses to a synthetic environmental variable generated by a principle components analysis. The best fit was obtained with a sigmoid function which explained between 74 and 93% of the variation in sap flux of individual trees. Sap flow reached an asymptote where higher light and evaporative demand did not cause sap flux to increase further. Soil moisture had little influence on sap flux. The morphological characteristics of the trees significantly affected sap flow; taller trees responded to changes in environmental variables sooner than shorter trees and high liana cover buffered tree sap flow responses to weather. The effect of species‐specific differences on the model was small; the mean effectiveness of the model was reduced by 6% when parameters were estimated from a single pool of measurements taken from all individuals. The results indicate that sap flow response could be effectively estimated using a simple general model and composite environmental index for these 10 diverse tree species.     

黄土丘陵区辽东栎群落优势种和主要伴生种树干液流动态特征

运用Granier热扩散探针法(TDP),于2011年5月对黄土丘陵区延安市南郊公路山辽东栎群落优势种(辽东栎)和3个伴生种(山杏、细裂槭、桃叶卫矛)树干液流进行连续测定,并同步监测气象环境因子(太阳辐射、空气温度和相对湿度)和土壤水分动态,比较分析辽东栎、山杏、细裂槭、桃叶卫矛的树干液流通量密度日变化动态及其对环境因子的响应特征。结果表明,辽东栎、山杏、细裂槭和桃叶卫矛液流通量密度日变化动态特征总体上反映了气象环境因子变化的昼夜规律性,呈单峰或双峰型曲线,但辽东栎与山杏、细裂槭、桃叶卫矛液流通量密度日变化规律存在一定的差异。辽东栎液流启动时间较早,通常在5:00左右,而山杏、细裂槭和桃叶卫矛液流启动时间均晚于辽东栎一个小时以上。辽东栎液流达到峰值的时间也较其它树种早,通常出现在9:00前后;细裂槭、桃叶卫矛和山杏液流通量密度达到峰值的时间分别在10:00、11:00和13:00前后。细裂槭和桃叶卫矛液流通量密度日变化曲线峰值较窄。除此之外,土壤水分状况对伴生种的影响程度要比优势种的大。相关分析表明,树干液流通量密度与太阳辐射和空气水汽压亏缺均呈极显著正相关。优势种与伴生种的液流动态差异可能与树木本身特性以及群落对光能的限制性再分配有关。     

疏伐对黄土丘陵区刺槐林蒸腾的影响

黄土高原地区植被建设已达到土壤水分承载力的阈值,需要对现有林分进行结构优化并提升其生态功能。不合理的林分密度是导致黄土丘陵区刺槐林土壤干化、生长衰退的主要原因之一。疏伐可以优化林分结构,并能够通过控制蒸腾耗水来调控土壤水分,是促进刺槐林可持续生长的有效手段。疏伐对黄土丘陵区刺槐林蒸腾有何影响,目前并不清楚。研究基于树干液流法估算了4个不同疏伐强度（样地1：52%、样地2：48%、样地3：35%、样地4：未疏伐）下刺槐单株尺度的液流速率与林分尺度的日平均蒸腾量,并分析了不同时间尺度下液流速率与环境因子的关系,以阐明疏伐对黄土丘陵区刺槐林蒸腾的影响。结果表明：（1） 单株尺度刺槐蒸腾速率（即液流速率）随疏伐强度减小（林分密度增大）呈现下降趋势（样地1：0.53 kg cm^-2 d^-1、样地2：0.41 kg cm^-2 d^-1、样地3：0.31 kg cm^-2 d^-1、样地4：0.33 kg cm^-2 d^-1）;（2） 观测期林分尺度日平均蒸腾量随疏伐强度减小呈现上升趋势（样地1：0.90 mm/d、样地2：1.18 mm/d、样地3：1.04 mm/d、样地4：1.44 mm/d）;（3） 在半小时尺度与日尺度上,各样地液流速率与环境因子的关系没有显著差异,半小时尺度单株液流速率均与太阳辐射相关性最高（相关系数0.883-0.908）,液流速率日变化过程与环境因子日变化过程存在时滞现象;日尺度单株液流速率与饱和水汽压亏缺相关性最高（相关系数0.843-0.913）,样地间日尺度单株液流速率的差异性随着饱和水汽压亏缺增大而增大。研究结果初步反映了疏伐导致的林分密度变化对刺槐蒸腾的影响,将为黄土丘陵区刺槐林的结构改造、功能提升和土壤水分调控提供理论支持。     

Calcium in Xylem Sap and the Regulation of its Delivery to the Shoot

Amounts of total and free calcium in root and shoot xylem sapwere quantified for a number of species grown in comparableenvironments and in a rooting medium not deficient in calcium.The potential for the shoot to sequester calcium was also examined,along with the ability for ABA to regulate calcium flux to theleaf. Xylem sap calcium showed considerable interspecific and diurnalvariation, even though the plants were grown with similar rhizosphericcalcium concentrations. The potential for the shoot to sequesterxylem sap calcium was also highly variable between species andimplied an ability, at least in some species, to regulate thecalcium reaching the shoot in the transpiration stream. Long distance transport of calcium in the xylem was not primarilyby mass flow, because neither calcium uptake nor distributionwere closely related to water uptake or transpiration. The diurnalchanges in xylem sap total ion concentration appeared to benegatively correlated with transpiration while, in contrast,the calcium ion concentration showed two peaks, one occurringin the dark and the other in the light period. The application of ABA to roots caused an increase in the rateof exudation from the xylem of detopped well-watered plants.These experiments suggest that changes in root water relationsdriven by ionic fluxes were the likely cause for enhanced sapexudation from ABA-treated roots. The steady-state concentrationof calcium in the xylem sap was unaffected by ABA when exudationrate increased and, consequently, the flux of calcium must alsohave increased. Key words: Abscisic acid, calcium, xylem sap, ionic fluxes     

胡杨液流对地下水埋深变化的响应

以塔里木河流域荒漠河岸林主要建群种胡杨(Populus euphratica)为研究对象, 结合中下游不同断面地下水埋深和胡杨液流变化的监测数据, 分析了胡杨茎干液流与地下水埋深变化的关系, 探讨荒漠环境下天然胡杨生长的合理生态水位。研究表明, 胡杨液流通量密度随地下水埋深即干旱胁迫程度的加大而减小, 两者呈极显著负相关, 相关系数达-0.887; 胡杨液流通量在地下水埋深位于4.5-5 m时出现异常变化, 表明此时胡杨的正常生长受到胁迫, 胡杨通过自身调节降低蒸腾耗水以适应环境; 土壤盐分不是影响塔里木河中下游各断面胡杨液流变化的主要因子; 对植物样地调查结果分析显示, 胡杨盖度、密度和频度均在地下水埋深在4-6 m梯度下开始表现为降低趋势。综合分析认为维系塔里木河中下游天然胡杨正常生长的生态水位为地下水埋深4.5 m以内。     

Xylem sap composition: A tool for investigating mineral uptake and cycling in adult spruce

Xylem sap composition of spruce is influenced by several factors, such as the sampled organ, the sampling period, the availability of soil nutrients, and the soil water potential. Based on literature data and ongoing investigations carried out with adult trees, we present an overview on the main factors influencing xylem sap concentrations of Norway spruce. Direct measurements of nutrient fluxes in the xylem sap are then used to suggest a general scheme of mineral element cycling within adult trees. In Norway spruce (Picea abies Karst.), nutrient concentration in the xylem sap was higher in twigs and fine roots compared to the bottom of the trunk, the highest concentrations beeing observed in spring during the shoot elongation. Xylem sap concentrations were higher in spruce growing at nutrient rich sites than at poor sites. The combination of twig and trunk xylem sap analysis, together with xylem flow measurements in the trunk during the course of a vegetation period allowed the quantification of mineral fluxes via xylem sap flow in the trunk and twigs. These results were compared to gross mineral uptake measurements at the same site. Ca flux in the trunk xylem sap was lower than the gross uptake of Ca. Mg flux in trunk sap was approximately equivalent to Mg gross uptake whereas P and K fluxes in trunk sap were much higher than the gross uptake. Fluxes of Ca, Mg, K and P in the twig sap were much higher than that in trunk sap. Data suggest that internal cycling is responsible for a large part of the nutrient fluxes in the xylem sap of the crown. Xylem sap composition thus appears to be a tool which can complement other sources of information on mineral uptake and cycling in adult spruce     

Seasonal and spatial variation of carbohydrates in mistletoes (Viscum album) and the xylem sap of its hosts (Populus x euamericana and Abies alba)

In the present field study we analysed the seasonal pattern of carbohydrate composition and contents in the xylem sap of Viscum album and the xylem sap of a deciduous ( Populus × euramericana ) and a coniferous ( Abies alba ) host tree species. The results were compared with the soluble carbohydrate composition and contents of mistletoe tissues. On both hosts significant amounts of glucose, fructose, and sucrose were found in the xylem sap of Viscum throughout the seasons. The general seasonal pattern of sugar contents, i.e. high concentrations in spring and lower concentrations in other seasons on Populus , and intermediate concentrations throughout the year on Abies , largely reflected the xylem sap carbohydrate composition and contents of the respective host. These observations provide indirect evidence for carbohydrate flux from the xylem sap of the host into the mistletoe. However, in both hosts xylem sap seems to be deviated into the mistletoe without specific control of carbohydrate flux. Differences observed between the seasonal pattern of xylem sap carbohydrate concentrations in Viscum on Populus and Abies may originate from the different time of leaf development of these species. A clear-cut seasonal pattern of soluble carbohydrates was not observed in the leaves of Viscum on both hosts. Still soluble carbohydrates seem to be reallocated from the senescing to the newly developed leaves of Viscum indicating that the seasonal requirement of carbohydrate for growth and development can only completely be met by carbohydrate acquisition from the host and their own photosynthesis.     

黄土丘陵区两典型造林树种生长季树干直径微变化动态及其影响因素

运用DC3型高分辨率树干直径变化记录仪和Granier热扩散探针,对黄土丘陵地区两典型树种辽东栎和刺槐生长季的树干直径微变化和树干液流动态进行连续监测,并同步观测主要环境因子(土壤含水量、太阳辐射、空气温度和相对湿度),分析两树种树干的直径微变化动态特征与蒸腾耗水的关联性及其对环境因子的响应。结果表明: 两树种树干直径和液流通量密度呈现明显的昼夜变化规律,直径日最大收缩量与日均液流通量密度呈显著正相关,树干直径在日尺度上的微变化受当日蒸腾耗水量的影响。对树干直径日最大收缩量和蒸腾驱动因子进行线性拟合,结果显示,树干直径日变化量与主要气象环境因子(日均太阳辐射、日均空气水汽压亏缺、整合变量VT)呈显著正相关。回归曲线斜率表明,辽东栎树干直径日变化量大于刺槐,其直径对气象环境因子的敏感度更大。两树种液流通量密度在较高土壤水分时段高于土壤含水量较低时段,在不同土壤水分条件下辽东栎树干直径日最大收缩量差异显著,刺槐未达到显著水平,这些差异可能与两树种蒸腾调节和树干水分补充等用水策略有关。     

Effect of Far Red Light on Root Growth and on Xylem Sap in Cotton (Gossypium hirsutum L.)

Far red light given before darkness highly increased droughtresistance of cotton plants. There was higher stomatal resistancein leaves resulting in a lower transpiration rate. The aim ofthis work was to examine the effect of far red light on rootgrowth and its effect on water transfer measured by analysisof xylem sap obtained after cutting the hypocotyl. Short-termand long-term treatments of far red light were used. In thepresence of far red light, the amount of xylem sap decreasedwhile root resistance increased. Qualitative analysis of thexylem sap indicated an increase of amino acid content and analteration of potassium flux. Calcium flux was not significantlymodified. The change in potassium content of xylem sap is discussedwith regard to turgor and drought resistance. (Received June 9, 1984; Accepted October 8, 1985)