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.     

Compensation heat-pulse measurements of sap flow for estimating transpiration in young lemon trees

Potted two-year-old lemon trees [Citrus limon (L.) Burm. f.], cv. Verna grafted on sour orange (C. aurantium L.) rootstock, growing in greenhouse, were subjected to drought for 33 d. Control plants were daily irrigated at field capacity. Values of sap flow (SF) were compared with transpiration (E) rates measured gravimetrically. The results underlined the robustness and high sensitivity of the compensation heat-pulse technique for estimating transpiration on a wide range of SF. Good direct correlations between E and SF rates on an instantaneous and daily basis were obtained in both treatments. On a daily basis, a common calibration curve can be used for both irrigation treatments. On an instantaneous basis, changes in SF were matches by similar changes in E in both treatments, although the relationships between these parameters presented different intercepts in each treatment. Sap flow rates were influenced by weather conditions in trees growing in non-limiting soil water conditions. This makes it possible to evaluate the significance of any sap flow measurement in relation to the reference value calculated for the vapour pressure deficit at the time the measurement was taken.This research was supported by Ministerio de Educacion y Ciencia (MEC), (CICYT/FEDER AGL2003-9387-C05-02 and AGL2004-0794-C03-02), and PETRI (PTR1995-0693-OP-02-01) grants to the authors. M.F. Ortuno was a recipient of a Program I3P research fellowship from CSIC.     

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.     

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.     

Changes in stem water content influence sap flux density measurements with thermal dissipation probes

Key message

Stem WC may decline during the day. Zero-flow dT _m increases when WC decreases. Use of nighttime dT _m in the calculation of sap flux density during the day might introduce errors.

Abstract

There is increasing evidence of diel variation in water content of stems of living trees as a result of changes in internal water reserves. The interplay between dynamic water storage and sap flow is of current interest, but the accuracy of measurement of both variables has come into question. Fluctuations in stem water content may induce inaccuracy in thermal-based measurements of sap flux density because wood thermal properties are dependent on water content. The most widely used thermal method for measuring sap flux density is the thermal dissipation probe (TDP) with continuous heating, which measures the influence of moving sap on the temperature difference between a heated needle and a reference needle vertically separated in the flow stream. The objective of our study was to investigate how diel fluctuations in water content could influence TDP measurements of sap flux density. We analysed the influence of water content on the zero-flow maximum temperature difference, dT _m, which is used as the reference for calculating sap flux density, and present results of a dehydration experiment on cut branch segments of American sycamore (Platanus occidentalis L.). We demonstrate both theoretically and experimentally that dT _m increases when stem water content declines. Because dT _m is measured at night when water content is high, this phenomenon could result in underestimations of sap flux density during the day when water content is lower. We conclude that diel dynamics in water content should be considered when TDP is used to measure sap flow.     

A new method of sap flow rate determination in trees

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

A requirement for sucrose in xylem sap flow from dormant maple trees

The response of excised stem segments of several tree species to freezing and thawing cycles was studied. All species studied (Thuja occidentalis, Fagus grandifolia, and Betula papyrifera) except maple (Acer spp.) exuded sap while freezing and absorbed on thawing. Maple stems absorbed sap while freezing and exuded sap during the thaw only when sucrose was present in the vessel solution. Increased concentration of sucrose in the vessel sap led to increased exudation. In the absence of sucrose, maple stems absorbed sap on thawing. The presence of sucrose enhanced sap absorption during freezing cycles in maples. In general, large sugars, disaccharides and larger, could substitute for sucrose in the maple exudation response while sugar hexoses could not. The results are discussed in relation to the O'Malley-Milburn model (1983 Can J Bot 61: 3100-3106) of sap flow in maples.     

Improvement of the heat pulse method for determining sap flow in trees

Abstract. The heat pulse method for determining sap flux in large woody sterns was modified for easier field operation. It uses the measurement of the time elapsed between heat pulse release by a line heater radially inserted in the stem, and the occurrence of maximum temperature 15 mm downstream of the heater. This spacing between heater and thermometer is critical to the reliability of the measurement. Calculations using uncorrected theory provide estimates of the sap flux density in stems with both uniform and non-uniform cross-sectional distribution of conducting tissues which are about 55% of the actual sap flux density. This factor results from insufficient thermal homogeneity between tissues where sap flow occurs and tissues where sap flow has been interrupted.
Sap flow in trunks of citrus trees was inferred from measurements of the cross-sectional distribution of sap flux density. Variability of sap flux density is specific to each trunk and is time-dependent and imposes multiple radial and angular measurements. The method was checked in a citrus trunk ramified into three branches. Instantaneous determinations of the flow in the trunk and in the branches differed by less than 5.7%. The daily values agreed within 2.8%.     

热消散液流测定系统研究竹子蒸腾的问题和解决思路

旺盛的蒸腾是竹子迅速生长的重要代谢保障,对林区生态系统水循环和水量平衡发挥关键作用,如何准确估测竹林的蒸腾耗水是迫切需要解决的方法难题。目前研究树木水分生理和森林水文广泛采用的热消散液流测定方法(TDP)却少有在竹林中应用,而仅有的几项研究报道由于缺乏严谨的实验验证,结果误差较大而缺少说服力。作者认为,竹壁结构异质性和竹竿中央空腔造成热量的不均匀分布是基于热通量交换的液流测定系统测量竹子蒸腾不准确的主要原因,提出以自行设计的注水变压液流特性测定法和传统的整树容器称重法,验证热消散液流测定系统估测竹子水分利用的适用性。本文还结合竹子的种群结构和无次生生长的特点,提出竹株借助相互连接的地下茎进行水分再分配,异株补偿水力限制,蒸腾主要受年龄而不是竹形大小影响的观点,建议通过分析竹子蒸腾的年龄效应,研究不同种植密度竹林水分利用的变化规律,为竹林生产和集水区水源管理提供有效的林型设计和措施。     

树木树液上升机理研究进展

水分在植物体内的运输一直是很多植物生理生态学家所关注的一个重要问题。介绍了内聚力学说的基本假设和其存在争议,总结了近年来这一研究领域的几个热点问题,主要包括：（1）木质部栓塞及其恢复机理;（2）木质部压力探针和压力室法测定的木质部张力值不一致的现象及其可能原因;（3）补偿压学说;（4）不同界面层张力以及输水管道的毛细作用力、薄壁细胞膨压和木质部渗透压、逆向蒸腾等在树木汁液上升中的贡献;（5）最近发现的存在于木质部导管伴胞和韧皮部薄壁细胞等质膜中的水孔蛋白在植物水分运输中的调控作用等。这些方面在解释树木的树液上升中都起着重要的作用。     

树干自然温度梯度变化对热扩散法测算树干液流速率的影响

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

Applications of a thermal imaging technique in the study of the ascent of sap in woody species

We present applications of infrared thermograph to the direct observation of water transport in stems of woody species. Presently, the method gives only qualitative information on the path of sap movement along the stem, and therefore, does not replace any of the common quantitative methods of sap flow measurement. Nevertheless, the thermal imaging technique provides a novel approach to the study of the ascent of sap and could have a role in supporting more quantitative methods. Thermography permits visualization of the thermal variations of a fairly large area of the stem in real time, and hence, is well suited for spatial analysis of sap movement. Three experiments were carried out during the summers of 1991 and 1992, with the following objectives: to define the sap flow distribution in the active annual rings; to identify grain anomalies in the sap wood; and to study the consequences of induced embolism on the path of sap flow. Altogether, we tested more than 10 woody species (broad-leaves and conifers) either in natural conditions or in the nursery. We found considerable differences in the spatial distribution of sap flow not only between species and individuals but also within a single tree. Grain anomalies or embolized zones in the xylem, which are easily demonstrated, often further modify the path of sap flow. The irregularity of sapwood dimensions and of sap flow among tree rings requires careful evaluation of the positioning of the probes when using methods for quantitative measurements.     

A comparison of sap flow measurements and potometry in two tropical lowland tree species with contrasting wood properties

We evaluated the performance of the Heat Dissipation Technique (HDT) to measure sap flow in whole trees by comparison with potometric water uptake. Two tropical lowland species, Ochroma lagopus (balsa), a pioneer species with light wood and Hyeronima alchorneoides (pilón), a late-successional species with hard wood were examined. Diurnal courses of sap flow measured with the HDT showed good agreement with potometry. At the low sap flow rates (below 1 Kg h(-1)) occurring during nocturnal recharge HDT consistently underestimated sap flow rates. This resulted in the failure of the current version of the HDT to measure nocturnal water uptake, an important component of the water budget of at least one of the two species examined.     

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.     

A method for calibration of flow cytometric wavelength shift fluorescence measurements

Recently, new fluorescent dyes have been introduced into flow cytometry which alter their spectral characteristics when changes occur in certain cell features, e.g., intracellular pH or calcium ion concentration. Such changes may be determined by measuring the fluorescence intensity ratio in two different wavelength ranges (5). Here a new method is described, which simplifies the use of steadily flowing fluids for calibration. The pulse electronics of a flow cytometer cannot process the static fluorescence signals of a streaming fluid. If, however, the exciting or emitted fluorescence light of a calibration fluid is made pulsating, the flow cytometer electronics can evaluate those pulses. The new calibration procedure uses measurement of two wavelength windows shown in a two-parameter display to generate an absolute calibration scale. Measurement of the spectral shift in calibration fluids under identical instrumental settings provides absolute values that measurements of intracellular concentrations can be referred to.     

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.     

A simple technique for thermal stimulation of single semicircular canals.

Two simple thermodes for stimulating single semicircular canals have been designed in order to elicit eye nystagmus. The first, for warm stimulation, is based on the Joule effect: a silver needle is warmed up by means of a coiled nickrome wire. The temperature at the thermode tip is function of current intensity. The thermode for cold stimulation is a device in which a circulating freezing mixture cools a silver needle. The temperature at the thermode tip is proportional to the temperature of the freezing mixture.