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.     

A gauge to measure the mass flow rate of water in trees

A gauge that measures the mass flow rate of water in a growing tree is described. The gauge consists of an electric band heater wrapped around a section of the stem, a temperature controller that switches the current to the heater on and off so as to maintain a constant temperature rise across the heated section, and a timer to record the total time for which the heater is switched on. An energy balance shows the mass flow of water to be proportional to the time of operation of the heater. The magnitude of measurement errors is estimated using dimensional analysis and a numerical model. Experimental measurements of the flow rate of water in a tree using the gauge agree well with its recorded loss of weight.     

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.     

The Relationship between Transpiration, Root Water Uptake, and Leaf Water Potential

The effect of changing the transpiration rate on leaf waterpotential and water balance has been examined to show if permeabilityof the plant (predominantly the roots) is constant or varieswith the transpiration rate. Measurements of leaf effectivethickness, water potential, transpiration, and uptake of waterby roots were made on sunflower, barley, and maize plants grownin solution culture and subjected to a range of atmosphericconditions and root treatments: cooling, low osmotic potential,and removal of part of the root system. Leaf water potential changed little under a wide range of atmosphericconditions and rates of water flux in the three species, sothat the root permeability to water increases as the rate oftranspiration, and therefore flow across the root surface, increases.Equality between uptake and loss of water and thereby maintenanceof constant leaf water potential is assisted by stomatal changes,which appear to be in response to conditions at or in the rootrather than a direct response to changes in bulk leaf waterpotential.     

Stomatal oscillations in orange trees under natural climatic conditions

BACKGROUND AND AIMS: Stomatal oscillations have been reported in many plant species, but they are usually induced by sudden step changes in the environment when plants are grown under constant conditions. This study shows that in navel orange trees (Citrus sinensis) pronounced stomatal oscillations occur and persist under natural climatic conditions. METHODS: Oscillations in stomatal conductance were measured, and related to simultaneous measurements of leaf water potential, and flow rate of sap in the stems of young, potted plants. Cycling was also observed in soil-grown, mature orchard trees, as indicated by sap flow in stem and branches. KEY RESULTS: Oscillations in stomatal conductance were caused by the rapid propagation and synchronization of changes in xylem water potential throughout the tree, without rapid changes in atmospheric conditions. CONCLUSIONS: The results show marked stomatal oscillations persisting under natural climatic conditions and underscore the need to discover why this phenomenon is so pronounced in orange trees.     

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     

Water Balance in Cucumis Plants, Measured by Nuclear Magnetic Resonance, I

In this paper we demonstrate the study of plant water balanceby the non-invasive measurement of tissue water content andwater flow using proton nuclear magnetic resonance (NMR). Sapvelocity and flux were measured independently in the presenceof an excess of stationary tissue water. The instrumentationdescribed allows automated and unattended measurement of flow-and water content-variables in a well-defined region of theplant over periods of several days, with a time resolution betweensuccessive measurements of c. 5 s. Using this apparatus theeffect of changes in light intensity (day/night rhythm) andrelative humidity on stem tissue water content as well as onthe velocity and flux of xylem sap in the stem were investigatedin a cucumber plant. The results are in agreement with predictionsfrom a simple model for plant water balance, which is basedon water potential, flow rate and resistance to flow. As longas only transpiration is varied, flow rate and water content(or potential) are affected in opposite ways as demonstratedin this paper. In contrast, the model predicts that changesin uptake (resulting from changes in, for example, root resistance)will induce changes in water content and flow in the same direction.An experimental verification of this prediction is given ina subsequent paper, where, in addition, the NMR results arecompared to those obtained with a dendrometer. Key words: Water balance model, Cucumis sativus L., flow, water content, NMR, water balance measurement     

Transpiration measurements on apple trees with an improved stem heat balance method

Since the late eighties a handy and user-friendly sap flow meter (Dynagage®) is on the market which can quantify 0205 the sap flow through intact plant stems, based on the stem heat balance method. The documentation about its accuracy and reliability, however, is still too limited to use it as a standard method in field experiments with apple trees. We therefore tested this commercial system on potted apple trees (Malus domestica L.; cv. Red Elstar and Jonagold; on rootstock M9 vf) with stem diameters of 1.8 to 4 cm. The measured sap flow was compared with mass loss measured by an automated balance, supposing the total mass loss of the trees was equal to the water loss by transpiration. The results revealed three major problems:

Artificially Induced Water and Sulfate Transport through Sunflower Roots

A new experimental method is used to determine simultaneously the quantity and composition of the sap exuded by a detopped root system at the same time that a pressure deficit of desired magnitude can be applied to the stem stump. The technique was used in a study of the transport of radioactive sulfate through the roots of young sunflower plants placed on complete nutrient solutions labelled with ³⁵S. The complications by the time factor on the composition and rate of the sap stream in experiments of this type were observed and discussed. The time of detopping the roots was very critical as the conditions of sulfate transport were greatly changed some time after the excision. A rectilinear connection existed between the rate of sulfate transport in the sap and the water flow at sap flow velocities comparable with transpiration rates. When the transport of water was very slow, the rate of sulfate transport became constant and independent of the water stream. It was suggested that diffusion or water flow could act as motive force for the ion transport in some non-metabolic phase of transfer in the roots. The addition of 2,4-DNP to the test solution severely interfered with the water and sulfate transport conditions in the roots.     

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。     

Coppicing affects growth, root:shoot relations and ecophysiology of potted Quercus rubra seedlings

Two experiments were conducted to examine the response of Quercus rubra L. seedlings to coppicing. In a greenhouse experiment, growth, biomass distribution, leaf gas exchange, and water and carbohydrate relations were measured for 1-year-old seedlings that were either coppiced when dormant at the time of planting or left intact as controls. Coppicing induced sprouting from the base of the stem, and, in general, the physiology of sprouts and controls was similar. However, the relative growth rate (RGR) of sprouts was 9% higher than that of controls, allowing sprouts to compensate fully for the initial mass lost to coppicing. In a second experiment, in an outdoor cold frame, growth, biomass distribution, leaf gas exchange and plant water relations were measured on 1-year-old seedlings that were either coppiced at the time of planting (dormant-coppiced), coppiced soon after bud break (active-coppiced) or left intact (controls). Dormant coppicing again had little impact on seedling physiology, and dormant-coppiced plants again compensated for initial mass loss with a higher RGR. In contrast, active-coppiced seedlings did not compensate for initial mass loss, as their RGR did not differ from that of controls. By the tenth week of the study, leaf gas exchange rates of active-coppiced sprouts were higher than those of dormant-coppiced and control seedlings. Active-coppiced sprouts also had a greater soil-to-leaf hydraulic conductivity (expressed on a leaf area basis) and a lower ratio of leaf area to root surface area than did controls. Across treatments, photosynthetic rate and stomatal conductance were positively correlated with soil-to-leaf hydraulic conductivity, and gas exchange rates and hydraulic conductivity were negatively related to leaf:root area ratio. Thus, the removal of actively growing shoots may have altered subsequent leaf gas exchange largely through coppice-induced changes in leaf-root balance.     

Effects of vegetation on flow through free water surface wetlands

The one-dimensional Saint-Venant equations are modified to account for stem drag and volumetric displacement effects of dense emergent plants on free surface flow. The modified equations are solved with an implicit finite difference method to give velocities and depths for shallow flows through a vegetated wetland channel. Estimated flow profiles are used to investigate how vegetation density, downstream boundaries and aspect ratio affect detention time, an important parameter in determining nutrient and pollutant removal efficiencies of wetlands constructed to treat wastewater. Results show that free water surface wetlands may exhibit static, neutral or dynamic behavior. Under static conditions, the wetland behaves like a pond in which displacement effects caused by submerged plant mass invariably decrease detention times. Under dynamic conditions, stem drag induced by aquatic plants predominates and wetland detention times increase with vegetation density. These opposing responses are separated by a narrow neutral condition where the presence of vegetation has virtually no net effect on detention time. For a given flow rate and surface area, detention times and hence treatment efficiencies in vegetated free water surface wetlands can be managed to some degree by adjusting the downstream control or by changing the aspect ratio.     

Quantification of water transport in plants with NMR imaging

A new nuclear magnetic resonance imaging (NMRi) method is described to calculate the characteristics of water transport in plant stems. Here, dynamic NMRi is used as a non-invasive technique to record the distribution of displacements of protons for each pixel in the NMR image. Using the NMR-signal of the stationary water in a reference tube for calibration, the following characteristics can be calculated per pixel without advance knowledge of the flow-profile in that pixel: the amount of stationary water, the amount of flowing water, the cross-sectional area of flow, the average linear flow velocity of the flowing water, and the volume flow. The accuracy of the method is demonstrated with a stem segment of a chrysanthemum flower by comparing the volume flow, measured with NMR, with the actual volumetric uptake, measured with a balance. NMR measurements corresponded to the balance uptake measurements with a rms error of 0.11 mg s(-1) in a range of 0 to 1.8 mg s(-1). Local changes in flow characteristics of individual voxels of a sample (e.g. intact plant) can be studied as a function of time and of any conceivable changes the sample experiences on a time-scale, longer than the measurement time of a complete set of pixel-propagators (17 min).     

Water relations of the tomato during fruit growth

Fruit and stem water potentials of tomato plants were measured continuously for several days using automated psychrometers. A linear voltage displacement transducer was used to simultaneously measure diameter changes on an adjacent fruit. A strong correlation was observed between the water potential gradient of the fruit and stem, and changes in fruit diameter. Fruit diameter increased when the apoplasmic water potential gradient favoured solution flow into the fruit and fruit shrinkage occurred only when the water potential gradient was inverted. Based on our data and other published data (Ehret & Ho 1986; Lee 1989a) on phloem transport in tomato, we have concluded that low stem water potentials have an immediate and direct effect on phloem turgor; reducing the driving force for sap flow into the fruit. Since fruit water potential remained relatively constant, the diurnal variation in stem water potential was sufficient to account for the correlation with changes in fruit diameter. There are consequences with respect to predicting the accumulation of dry matter in tomato fruit.     

Water use of a multigenotype poplar short‐rotation coppice from tree to stand scale

Short‐rotation coppice (SRC) has great potential for supplying biomass‐based heat and energy, but little is known about SRC's ecological footprint, particularly its impact on the water cycle. To this end, we quantified the water use of a commercial scale poplar (Populus) SRC plantation in East Flanders (Belgium) at tree and stand level, focusing primarily on the transpiration component. First, we used the AquaCrop model and eddy covariance flux data to analyse the different components of the stand‐level water balance for one entire growing season. Transpiration represented 59% of evapotranspiration (ET) at stand scale over the whole year. Measured ET and modelled ET were lower as compared to the ET of reference grassland, suggesting that the SRC only used a limited amount of water. Secondly, we compared leaf area scaled and sapwood area scaled sap flow (F_s) measurements on individual plants vs. stand scale eddy covariance flux data during a 39‐day intensive field campaign in late summer 2011. Daily stem diameter variation (?D) was monitored simultaneously with F_s to understand water use strategies for three poplar genotypes. Canopy transpiration based on sapwood area or leaf area scaling was 43.5 and 50.3 mm, respectively, and accounted for 74%, respectively, 86%, of total ecosystem ET measured during the intensive field campaign. Besides differences in growth, the significant intergenotypic differences in daily ?D (due to stem shrinkage and swelling) suggested different water use strategies among the three genotypes which were confirmed by the sap flow measurements. Future studies on the prediction of SRC water use, or efforts to enhance the biomass yield of SRC genotypes, should consider intergenotypic differences in transpiration water losses at tree level as well as the SRC water balance at stand level.     

开花期枣树茎流与气象因子的关系

阿克苏地区农田灌溉水资源利用率相对较低,大多采用漫灌为主、滴灌为辅,加上环境干旱、水资源紧缺已成为制约当地红枣种植业发展的主要因素,因此迫切需要寻找新型节水灌溉方法。该研究采用美国Dynamax公司生产的Flow-32包裹式茎流计对茎流速率进行连续监测,并在样地内安装HOBO小气候仪同步获取气象数据,将所得数据进行对比分析,探讨了不同灌溉方式下枣树茎流速率的变化规律以及与气象因子之间的关系。结果表明：两种灌溉方式下茎流连日变化呈现明显的昼夜变化规律,茎流速率均为单峰曲线且夜间茎流活动现象明显,早晨茎流启动时间W1早于W20.5 h,峰值前后W1和W2枣树茎流均出现较小幅度的“峰值群”,W1和 W2峰值出现时间分别为13：30－14：30和13：00－14：30,峰值大小分别为2133.33~2724.93 g·h-1和2234.11~2689.40 g·h-1;茎流日累积量变化过程曲线均为“S”形,但W1日茎流总量大于W2;茎流速率变化曲线均与太阳辐射和温度呈现正相关,与空气湿度呈现负相关,与风速相关性较差;太阳辐射表观波形变化趋势与两种灌溉方式下茎流变化曲线均存在异同,启动时间早于W1约0.5 h,W2约1 h,但峰值出现时间却晚于W1和W2。通过相关分析,太阳辐射均为主要影响因子,建立两种灌溉方式下液流速率与气象因子的多元线性模型,经过回归系数和相关系数检验,W1和W2多元线性回归方程均达到了显著水平。该研究结果为新型灌溉方式W1的实用性和优越性提供了科学依据。     

Evidence from Amazonian forests is consistent with isohydric control of leaf water potential

Climate modelling studies predict that the rain forests of the Eastern Amazon basin are likely to experience reductions in rainfall of up to 50% over the next 50-100 years. Efforts to predict the effects of changing climate, especially drought stress, on forest gas exchange are currently limited by uncertainty about the mechanism that controls stomatal closure in response to low soil moisture. At a through-fall exclusion experiment in Eastern Amazonia where water was experimentally excluded from the soil, we tested the hypothesis that plants are isohydric, that is, when water is scarce, the stomata act to prevent leaf water potential from dropping below a critical threshold level. We made diurnal measurements of leaf water potential (psi 1), stomatal conductance (g(s)), sap flow and stem water potential (psi stem) in the wet and dry seasons. We compared the data with the predictions of the soil-plant-atmosphere (SPA) model, which embeds the isohydric hypothesis within its stomatal conductance algorithm. The model inputs for meteorology, leaf area index (LAI), soil water potential and soil-to-leaf hydraulic resistance (R) were altered between seasons in accordance with measured values. No optimization parameters were used to adjust the model. This 'mechanistic' model of stomatal function was able to explain the individual tree-level seasonal changes in water relations (r2 = 0.85, 0.90 and 0.58 for psi 1, sap flow and g(s), respectively). The model indicated that the measured increase in R was the dominant cause of restricted water use during the dry season, resulting in a modelled restriction of sap flow four times greater than that caused by reduced soil water potential. Higher resistance during the dry season resulted from an increase in below-ground resistance (including root and soil-to-root resistance) to water flow.     

Water Relations of Sunflower (Helianthus annuus) Shoots during Exposure of the Root

The influence of root anoxia on the water balance of hydroponicallygrown sunflower plants was investigated in a controlled environmentroom. Leaf water potentials declined within 1 h of the onsetof anaerobic treatment of the roots. The severity of the waterstress during the initial 8 h of treatment was dependent onthe ambient humidity of the aerial environment. Recovery ofshoot water balance occurred by the beginning of the secondfull day of treatment, without stomatal closure. After 3 d of continuous anaerobic treatment, plant responsein the same experiment followed one of two patterns. In thefirst, there were declines in leaf water content, leaf waterpotentials, lamina expansion, transpiration rates and stomatalconductance. Osmotic adjustment occurred but it was not sufficientto prevent loss of turgor and the plants wilted at day 4 oftreatment. Complete rehydration took place after 6 d of treatmentif the stem was severed (under water) just below the cotyledons,indicating that shoot dehydration was a result of xylem occlusionat the root/shoot interface. Sections of the stem in this regionshowed xylem vessels to be discoloured and plate-like structuresobstructing the lumen. In the other pattern of response, plants did not develop a morenegative leaf water potential (except for an initial periodimmediately following treatment initiation) throughout the 6-danaerobic treatment period. There were no signs of water deficitalthough other symptoms such as precocious leaf senescence andepinasty occurred concomitantly with those in the first groupof plants that developed long-term water stress. Changes in shoot water relations are discussed in terms of ourearlier observations on anoxia-induced changes in the hydraulicconductivity and selectivity of the roots. Possible explanationsare offered for inter-plant variations in response to root anoxia. Key words: Anaerobiosis, water relations, xylem blockage     

Interrelationships of petiolar air canal architecture,water depth,and convective air flow in Nymphaea odorata (Nymphaeaceae)

• Premise of the study: Nymphaea odorata grows in water up to 2 m deep, producing fewer larger leaves in deeper water. This species has a convective flow system that moves gases from younger leaves through submerged parts to older leaves, aerating submerged parts. Petiolar air canals are the convective flow pathways. This study describes the structure of these canals, how this structure varies with water depth, and models how convective flow varies with depth. • Methods: Nymphaea odorata plants were grown at water depths from 30 to 90 cm. Lamina area, petiolar cross-sectional area, and number and area of air canals were measured. Field-collected leaves and leaves from juvenile plants were analyzed similarly. Using these data and data from the literature, we modeled how convective flow changes with water depth. • Key results: Petioles of N. odorata produce two central pairs of air canals; additional pairs are added peripherally, and succeeding pairs are smaller. The first three pairs account for 96% of air canal area. Air canals form 24% of petiolar cross-sectional area. Petiolar and air canal cross-sectional areas increase with water depth. Petiolar area scales with lamina area, but the slope of this relationship is lower in 90 cm water than at shallower depths. In our model, the rate of convective flow varied with depth and with the balance of influx to efflux leaves. • Conclusions: Air canals in N. odorata petioles increase in size and number in deeper water but at a decreasing amount in relation to lamina area. Convective flow also depends on the number of influx to efflux laminae.     

Gas-exchange and sap flow measurements of Salix viminalis trees in short-rotation forest

Simultaneous field measurements of transpiration and sap flow were performed on short-rotation Salix viminalis trees ranging in diameter from 1.5 to 3.5 cm (2-year-old shoots on 8-year-old stumps). Transpiration was measured using an open-top ventilated chamber enclosing the whole foliage of a tree. Sap flow was measured using a tree-trunk heat balance (THB) technique with a constant temperature difference and variable heat input. Both the instantaneous and daily values of water flux measured by the two absolute techniques agreed well with a difference of up to about 5%. In July, the hourly transpiration reached a maximum of about 0.2 kg m^–2 (leaf area) or 0.45 kg tree^–1, whereas maximum daily integrals reached 4 kg tree^–1. The response of sap flow rate to abrupt flux change when inducing emboli by cutting-off the stem was very rapid: the registered signal dropped by 85% within 10 min for a specimen with a projected leaf area of 2 m². For S. viminalis trees, transpiration was linearly correlated with stem cross-sectional area and with leaf area.