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     

新疆阿克苏干旱区富士苹果树干液流动态变化研究

选择新疆阿克苏干旱区富士苹果为研究对象,利用TDP茎流计连续测定苹果树干液流,并用自动气象站同步记录环境因子变化,探讨环境因子对树干液流的影响。结果显示:液流速率连日的变化过程是一个最大值不同的正弦曲线,夜间液流速率降低,白天液流速率上升;晴天液流速率呈明显的单峰曲线,阴天液流速率表现为多峰或双峰曲线,不论是晴天还是阴天,苹果在夜间仍然有微弱的活动,这种现象可能与新疆阿克苏地区的地理位置和气候有关;在苹果主要生长季内,树干液流速率(Fs)与大气温度(Ta)、太阳辐射(Rn)、相对湿度(RH)有较好的复相关关系(R2=0.93*),且影响苹果树液流速率Fs的最主要气象因子为大气温度Ta;苹果生长季内耗水总量为4 059.64L,并以7月耗水量最大(765.34L),占耗水总量的18.85%。研究表明,TDP径流计能够精确测定生长季内苹果茎干的液流速率与耗水量,且在干旱区影响苹果茎干液流变化的主要环境因子依次是大气温度、太阳辐射、相对湿度。     

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.     

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.     

油松、栓皮栎树干液流速率比较

应用TDP(ThermalDissipationProbe)技术对油松和栓皮栎树干液流进行了初步研究,经过野外近1a的实地定位观测,研究结果显示:栓皮栎月平均树干液流速率在整个生长期都较油松的月平均树干液流速率要高。前者大约是后者的5~10倍。栓皮栎在土壤干旱时期能够在白天产生明显的树干液流。在土壤干旱时期油松白天不产生树干液流而在晚上产生明显树干液流。在土壤相对湿润时期,油松和栓皮栎树干液流速率的波形与太阳总辐射的波形变化一致,但不同的是油松的树干液流速率波形呈明显的单峰状,而栓皮栎树干液流速率波形呈明显的多峰状。在土壤相对湿润时期太阳总辐射很低时能对油松树干液流速率产生明显的降低作用,而对栓皮栎树干液流则没有明显影响。在土壤干旱时期,油松和栓皮栎树干液流速率的峰值分别大约为0.0001cm/s和0.0006cm/s左右;在土壤水分充足时期,油松和栓皮栎树干液流速率的峰值分别大约相等约为0.0015cm/s左右,分别是油松和栓皮栎在干旱日期的液流速率峰值的10倍和2.5倍。     

CO2 efflux, CO2 concentration and photosynthetic refixation in stems of Eucalyptus globulus (Labill.)

In spite of the importance of respiration in forest carbon budgets,the mechanisms by which physiological factors control stem respirationare unclear. An experiment was set up in a Eucalyptus globulusplantation in central Portugal with monoculture stands of 5-year-oldand 10-year-old trees. CO₂ efflux from stems under shaded andunshaded conditions, as well as the concentration of CO₂ dissolvedin sap [CO₂^*], stem temperature, and sap flow were measuredwith the objective of improving our understanding of the factorscontrolling CO₂ release from stems of E. globulus. CO₂ effluxwas consistently higher in 5-year-old, compared with 10-year-old,stems, averaging 3.4 versus 1.3 µmol m^–2 s^–1,respectively. Temperature and [CO₂^*] both had important, andsimilar, influences on the rate of CO₂ efflux from the stems,but neither explained the difference in the magnitude of CO₂efflux between trees of different age and size. No relationshipwas found between efflux and sap flow, and efflux was independentof tree volume, suggesting the presence of substantial barriersto the diffusion of CO₂ from the xylem to the atmosphere inthis species. The rate of corticular photosynthesis was thesame in trees of both ages and only reduced CO₂ efflux by 7%,probably due to the low irradiance at the stem surface belowthe canopy. The younger trees were growing at a much fasterrate than the older trees. The difference between CO₂ effluxfrom the younger and older stems appears to have resulted froma difference in growth respiration rather than a differencein the rate of diffusion of xylem-transported CO₂. Key words: Eucalyptus globulus, refixation, stem respiration Received 19 May 2008; Revised 14 September 2008 Accepted 8 October 2008     

Thermal dissipation probe measurements of sap flow in the xylem of trees documenting dynamic relations to variable transpiration given by instantaneous weather changes and the activities of a mistletoe xylem parasite

The thermal dissipation probe was described in the early 1930s for the demonstration of a volume and mass flow of sap in the conductive elements of the xylem in trees. It was subsequently developed further and is now widely used in physiological ecology including measurements in the field. Thermal dissipation demonstrates the occurrence of sap flow and allows determination of its velocity. Here we report simultaneous continuous measurements of sap flow using the thermal dissipation technique and of transpiration by infrared gas analysis for diurnal and annual cycles in a deciduous and an evergreen oak tree, Quercus robur L. and Quercus turneri Willd., respectively, in a deciduous and an evergreen conifer, Larix decidua Mill. and Pinus griffithii McClell., respectively, and the host/mistletoe consortium of the deciduous linden Tilia mandschurica Rupr. & Max. and the evergreen Viscum album L. We show (1) that in diurnal cycles sap flow closely follows dynamic changes of the rate of transpiration elicited by daily fluctuations of weather parameters (sunshine, cloudiness, air temperature and humidity), (2) that in annual cycles sap flow reflects autumnal yellowing and shedding of leaves of the deciduous trees. We report for the first time comparative measurements of sap flow towards mistletoe shoots and host branches in a parasite/host consortium. This demonstrates (3) that mistletoes maintain considerably larger sap flow rates in their xylem conduits than the adjacent host branches dragging the transpiration stream of their host towards their own shoots. We also show (4) that even after the deciduous host has shed its leaves and itself does not transpire any more the evergreen mistletoe towards its shoots can maintain the persistence of a continuous sap flow via the stem and branches of the host as long as frost does not prevent that. The work presented underlines the contention that transpiration is the driving force for sap flow with continuous files of water in the xylem. It shows for the first time that mistletoes direct the flow of water through host roots and stems towards its own shoots by not only performing stronger transpiration as it is known from the literature but also by maintaining larger sap flow rates in the xylem conduits of its stems.     

Xylem sap proteins

Xylem sap from apple (Malus domestica Borkh), peach (Prunus persica Batsch), and pear (Pyrus communis L.) twigs was collected by means of pressure extrusion. This sap contained a number of acidic peroxidases and other proteins. Two other sources of xylem sap used in this study were stem exudates and guttation fluid. Similar peroxidases were also found in stem exudates and guttation fluids of strawberry (Fragaria x ananassa Duch.), tomato (Lycopersicum esculentum L.), and cucumber (Cucumis sativus L.). Isoelectric focusing activity gels showed that two peroxidases (isoelectric point [pl] 9 and pl 4.6) were present in initial stem exudates collected in the first 30 minutes after excision. Subsequent samples of stem exudate collected contained only the pl 4.6 isozyme. The pl 4.6 peroxidase isozyme was also found in root tissue and guttation fluid. These observations suggest that roots produce and secrete the pl 4.6 peroxidase into xylem sap. Cucumber seedlings were treated with 100 microliters per liter ethylene for 16 hours and the exudate from decapitated hypocotyl stumps was collected over a 3 hour period. Ethylene increased the peroxidase activity of stem exudates and inhibited the amount of exudate released. These observations suggest that xylem sap peroxidase may play a role in plugging damaged vascular tissue.     

鼎湖山针阔叶混交林优势种树干液流特征及其与环境因子的关系

运用Granier 热扩散式探针法,于2010年干湿季对鼎湖山自然保护区针阔混交林4种优势树种马尾松、锥栗、木荷和广东润楠的树干液流密度进行连续监测,并同步观测气温、相对湿度和光合有效辐射等环境因子的变化,研究其树干液流特征及其对环境因子的响应.结果表明： 在干湿季,4种优势树种的树干液流速率日变化均呈“昼高夜低”的典型单峰曲线,阔叶树锥栗、木荷和广东润楠的平均液流速率和峰值以及日液流量均显著大于针叶树马尾松;马尾松、锥栗、木荷和广东润楠的最大树干液流密度分别为29.48、38.54、51.67、58.32 g H₂O·m-2·s-1.优势树种树干液流速率的变化与环境因子的昼夜变化存在时滞;液流速率变化与光合有效辐射、水汽压亏缺和气温等环境因子的变化呈显著正相关,其中湿季以光合有效辐射为主导因子,干季以气温为主导因子.     

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.     

Time constant for water transport in loblolly pine trees estimated from time series of evaporative demand and stem sapflow

 The use of stem sap flow data to estimate diurnal whole-tree transpiration and canopy stomatal conductance depends critically upon knowledge of the time lag between transpiration and water flux through the stem. In this study, the time constant for water movement in stems of 12-year-old Pinus taeda L. individuals was estimated from analysis of time series data of stem water flux and canopy transpiration computed from mean daytime canopy conductance, and diurnal vapor pressure deficit and solar radiation measurements. Water uptake through stems was measured using a constant-heat sapflow probe. Canopy transpiration was correlated to stem uptake using a resistance-capacitance equation that incorporates a time constant parameter. A least-squares auto-regression determined the parameters of the resistance-capacitance equation. The time constants for ten loblolly pine trees averaged 48.0 (SE = 2.0) min and the time lag for the diurnal frequency averaged 47.0 (SE = 2.0) min. A direct-cross correlation analysis between canopy transpiration and sap flow time series showed maximum correlation at an approximately 30 min lag. Residuals (model-predicted minus actual stem flow data) increased with increasing soil moisture depletion. While the time constants did not vary significantly within the range of tree sizes studied, hydraulic resistance and capacitance terms were individually dependent on stem cross-sectional area: capacitance increased and resistance decreased with stem volume. This result may indicate an inverse adjustment of resistance and capacitance to maintain a similar time constant over the range of tree sizes studied.     

鼎湖山针阔叶混交林优势种树干液流特征及其与环境因子的关系

运用Granier热扩散式探针法,于2010年干湿季对鼎湖山自然保护区针阔混交林4种优势树种马尾松、锥栗、木荷和广东润楠的树干液流密度进行连续监测,并同步观测气温、相对湿度和光合有效辐射等环境因子的变化,研究其树干液流特征及其对环境因子的响应.结果表明:在干湿季,4种优势树种的树干液流速率日变化均呈“昼高夜低”的典型单峰曲线,阔叶树锥栗、木荷和广东润楠的平均液流速率和峰值以及日液流量均显著大于针叶树马尾松;马尾松、锥栗、木荷和广东润楠的最大树干液流密度分别为29.48、38.54、51.67、58.32g H2O·m-2·s-1.优势树种树干液流速率的变化与环境因子的昼夜变化存在时滞;液流速率变化与光合有效辐射、水汽压亏缺和气温等环境因子的变化呈显著正相关,其中湿季以光合有效辐射为主导因子,干季以气温为主导因子.     

Extraction of tracheal sap from Citrus and analysis of its nitrogenous compounds

Tracheal sap was extracted from sections of stems (0.5 to 1.5 cm in diameter and 7.5 to 15.0 cm in length) of orange trees (Citrus sinensis (L.) Osbeck cv. Washington Navel) by using a combination of the vacuum and liquid displacement methods. The volume of sap obtained and its concentration of nitrogenous compounds were dependent on the volume of displacing liquid used for the extraction. Four ml of water-saturated 1-butanol extracted essentially all of the xylem fluid present in the stem sections without apparent production of artifacts. The time of sampling affected the nitrogen concentration of the tracheal sap, but not the content of xylem nitrogen per volume of stem material. The orientation of the stems in the tree and the diameter of the stems had an effect on their contents of xylem nitrogen, with southeastern orientation and thinner stems showing higher concentrations. We could not detect the presence of ammonium, nitrites or proteins in the tracheal sap of orange trees. Most of the nitrogen was present as amino acids and about 2% of the total in the form of nitrates. The qualitative composition of amino acids, as determined by TLC, was the same both in winter and spring tracheal sap. The main components of the sap were proline and arginine in winter, and these amino acids together with asparagine and aspartic acid in spring.     

陇东旱地果园覆沙对苹果树蒸腾耗水及果实品质的影响

针对甘肃陇东旱地苹果园季节性干旱问题,以15年生长富2号苹果树为试材,测定果园覆沙后土壤水分、温度,以及果树主干液流速率、叶片气孔导度、果实品质等指标.结果表明：果园覆盖5 cm厚河沙,2-4月地温升高幅度低于1 ℃,6、7月晴天地温升高2.44 ℃,阴天地温升高2.61 ℃;在果树生长季节土壤含水量始终保持在田间持水量的60%以上.土壤含水量较高的时期（H期）晴天,树干液流曲线呈“几”字形宽峰曲线,覆沙处理液流启动时间较对照提前0.6 h,峰值较对照高25.5%,阴天峰值较对照高165.6%,且液流活动时间延长;土壤含水量较低的时期（L期）晴天,覆沙处理液流呈单峰曲线,启动时间较对照提前0.5～1 h,峰值在794 g·h^-1左右,而阴天液流启动时间较对照提前近1 h,峰值较对照高311.0%.3-7月对照的棵间蒸发量（E_s）较覆沙处理高156.0%,过多的地面蒸发是造成果树缺水的主要原因.覆沙后果实单果质量显著提高,果实硬度略有降低,果实可溶性固形物、维生素C、总糖、有机酸含量均有所提高.     

Composition of Amino Compounds Transported in Xylem of Casuarina sp.

Seedlings (180-d-old) of Casuarina cunninghamianaM L., C. equisetifoliaMiq. and C. glauca Sieber inoculated with each of two differentsources of Frankia, were analysed for translocated nitrogenouscompounds in xylem sap. Analyses were also made on sap fromnodulated and non-nodulated plants of C. glauca grown with orwithout a range of levels of combined nitrogen. Xylem exudateswere collected from stems, roots, and individual nodules ofnodulated plants and from stems and roots of non-nodulated plants.While the proportional composition of solutes varied, the samerange of amino compounds was found in xylem sap from the threedifferent symbioses. In C. glauca asparagine was the major aminoacid in the root sap followed by proline, while in symbioticC. cunninghamiana arginine accounted for more than 25% of theamino compounds. Citrulline was the major translocated productfound in the stem exudate of symbiotic C. equisetifolia. Increasingconcentrations of ammonium nitrate in the nutrient solutionresulted in increasing levels of free ammonia and glutaminein xylem sap from stems of nodulated and non-nodulated C. glauca,but there was relatively little change in the prominent solutes,e.g. citrulline, proline, and arginine. The composition of nitrogenoussolutes in stem or root exudates of C. glauca was similar tothat of exudate collected from individual nodules and on thisbasis it was not possible to distinguish specific products ofcurrent N₂ fixation in xylem. The main differences in N solutecomposition between the symbioses were apparently due to hostplant effects rather than nodulation or the levels of combinedN. Also, the data indicate that the use of the proportion ofN in sap as citrulline (or indeed any other organic N solute)could not be used as an index of nitrogen fixation.     

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。     

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     

Using branch and basal trunk sap flow measurements to estimate whole-plant water capacitance: a caution

Thermometric sap flow sensors are widely used to measure water flow in roots, stems and branches of plants. Comparison of the timing of flow in branches and stems has been used to estimate water capacitance of large trees. We review studies of sap flow in branches and present our own data to show that there is wide variation in the patterns and timing of sap flow of branches in different parts of the crown, owing to the course of daily solar illuminance. In contiguous forest, east-facing and upper branches are illuminated earlier than west-facing and lower branches and most capacitance studies do not include adequate information about branch sampling regimes relative to the overall pattern of crown illuminance, raising questions about the accuracy of capacitance estimates. Measuring only upper branches and normalising these results to represent the entire crown is dangerous because flows at the stem base likely peak in response to maximum crown illuminance (and transpiration) and this will differ compared to the timing of peak flows in upper branches. We suggest that the magnitude of flow lags between branches and stems needs further study, with careful attention to branch position and method application before a robust understanding of capacitance, particularly in woody tissues of large trees, can be formed. We did not detect flow lags in the world’s tallest and largest tree species Sequoia sempervirens and Sequoiadendron giganteum, despite measurement along large pathlengths (∼57 and 85 m), which raises questions as to why large flow lags are often recorded for much smaller species. One conspicuous possibility is the different methods used among studies. Constant-heating methods such as the thermal dissipation probe (and also heat balance methods) include heat capacitance behaviour due to warming of wood tissues, which delays the response of the sensors to changing sap flow conditions. We argue that methods with intrinsic heat-capacitance present dangers when trying to measure water-capacitance in trees. In this respect heat pulse methods hold an advantage.     

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.     

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%.