Sap feeding by the marsupial Petaurus australis: an enigmatic behaviour?

Summary The Yellow-bellied Glider, Petaurus australis, was observed to feed sporadically but extensively on eucalypt sap. Gliders extracted sap by making incisions into the bark with their procumbent lower incisor teeth and licking the resulting exudate. Less than 1% of possible trees were incised for sap. Sap flow and sap-sugar concentration were estimated for eight sap-site trees and eight non sap-site trees at intervals during an 18 month period. Measurements of sap-sugar concentration differed substantially among trees but fluctuated between sample periods independent of glider use. Sap flow measurements, were always higher for those trees currently in use by gliders, although the particular sap-site trees comprising this group varied between sample periods. The variation in sap flow occurred indenpendent of the incidence of rainfall or the type of microhabitat the tree occupied. Experimental incising of trees, designed to mimic the effects of feeding by gliders, failed to show any effect on sap flow. This suggests that the incidence of sap feeding is determined by a tree's pattern of sap flow and that there may be trees with unusual patterns of sap flow which gliders select as the most favourable trees to incise.     

The foraging behaviour of a nectar feeding marsupial,Petaurus australis

Summary The foraging behaviour of non-flying nectar feeding mammals has been examined rarely. The exudivorous yellow-bellied glider (Petaurus australis) was observed to feed extensively (70% of the total feeding observation time) on the nectar of all species of Eucalyptus present at a site in southeastern Australia. Gliders harvested nectar, and presumably pollen also, whenever eucalypt flowers were available and selected trees with 2–3 times as many flowers as that on trees randomly selected along a transect. The abundance of flowering trees varied temporally and, at times when few flowering trees were present, gliders chose trees with fewer flowers than at times when flowering trees were abundant. When flowering trees were superabundant or scarce, there was no relationship between the number of flowers in a tree and the duration of visits by gliders. However, at intermediate levels of abundance, the amount of time a glider spent in a tree was related to the number of flowers in a tree. Gliders devoted 90% of the time outside their dens to foraging and the above relationship is suggested to reflect two foraging options which maximize net energy gain for different abundances of flowering trees. Although gliders spent considerable lengths of time in individual trees feeding, initial deposition of cross pollen when gliders first arrive in a tree may be substantial and thus, may provide significant amounts of outcrossing for these eucalypts.     

Transpiration and stomatal behaviour of Quercus ilex plants during the summer in a Mediterranean carbon dioxide spring

Variations in the water relations and stomatal response of Quercus ilex were analysed under field conditions by comparing trees at two locations in a Mediterranean environment during two consecutive summers (1993 and 1994). We used the heat-pulse velocity technique to estimate transpirational water use of trees during a 5 month period from June to November 1994. At the end of sap flow measurements, the trees were harvested, and the foliage and sapwood area measured. A distinct environmental gradient exists between the two sites with higher atmospheric CO₂ concentrations in the proximity of a natural CO₂ spring. Trees at the spring site have been growing for generations in elevated atmospheric CO₂ concentrations. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits on water relations and whole-plant transpiration during the summer drought were severe. Leaf conductance and water potential recovered after major rainfall in September to predrought values. Sap flow, leaf conductance and predawn water potential decreased in parallel with increases in hydraulic resistance, reaching a minimum in mid-summer. These relationships are in agreement with the hypothesis of the stomatal control of transpiration to prevent desiccation damage but also to avoid ‘runaway embolism’. Trees at the CO₂ spring underwent less reduction in hydraulic resistance for a given value of predawn water potential. The decrease in leaf conductance caused by elevated CO₂ was limited and tended to be less at high than at low atmospheric vapour pressure deficit. Mean (and diurnal) sap flux were consistently higher in the control site trees than in the CO₂ spring trees. The degree of reduction in water use between the two sites varied among the summer periods. The control site trees had consistently higher sap flow at corresponding values of either sapwood cross-sectional area or foliage area. Larger trees displayed smaller differences than smaller trees, between the control and the CO₂ spring trees. A strong association between foliage area and sapwood cross-sectional area was found in both the control and the CO₂ spring trees, the latter supporting a smaller foliage area at the corresponding sapwood stem cross-sectional area. The specific leaf area (SLA) of the foliage was not influenced by site. The results are discussed in terms of the effects of elevated CO₂ on plant water use at the organ and whole-tree scale.     

Does a sap feeding marsupial choose trees with specific chemical characteristics?

Only about 20 vertebrate species are known to feed regularly on tree sap. One of these is the yellow‐bellied glider (Petaurus australis), a marsupial that obtains between 16% and 80% of its diet from eucalypt sap. We reviewed the literature on sap feeding by the yellow‐bellied glider and identified 10 species from which it most frequently obtains sap. These species come mainly from a few sections of the subgenus Symphyomyrtus, which accounts for two‐thirds of over 2000 sap trees reported, but more specifically sections Latoangulatae and Maidenii within this subgenus. Some of these key species contain relatively high concentrations of foliar nutrients and each is an important food tree for at least one of the arboreal folivores – koala (Phascolarctos cinereus), greater glider (Petauroides volans), and common ringtail (Pseudocheirus peregrinus) and brushtail possums (Trichosurus vulpecula). Because yellow‐bellied gliders, like the arboreal folivores, prefer to feed from certain individuals within a species, we hypothesized that these trees possess a unique chemical signature that links sap‐feeding with the concentrations of available nitrogen and formylated phloroglucinol compounds in leaves – the nutritive and defensive chemicals that influence feeding on these species by marsupial folivores. We tested this hypothesis in samples of leaves and bark collected from E. punctata and E. viminalis but found no link between chemistry and sap feeding and conclude that other aspects of an individual tree, such as sap flow or sap chemistry, determine whether gliders will target it for sap.     

Scots pine root distribution derived from radial sap flow patterns in stems of large leaning trees

This study characterizes whole tree root system distribution in a non-destructive way based on its functional parameters, particularly the sap flow patterns in stems. This approach particularly considers sap flow variation across stems, both radial and circumferential patterns of flow that are usually used for a better integration of sap flow density at the whole tree level. We focused at: (1) Showing examples of sap flow variation across stems at a defined situation (high midday values at the period of non-limiting water supply; (2) Analyzing radial flow patterns in terms of root distribution; (3) Validating these results at the stand level (mean data of series of individual trees) using results of classical biometric methods used at the same site; and (4) Applying the results for evaluation of root distribution around leaning trees. Sap flow rate was measured by the heat deformation method on a set of 14 trees at an experimental pine forest stand in Brasschaat (Belgium) during the growing season of 2000. Sap flow variation across stems was measured at a total of 700 points. Amounts of water supplied by superficial (horizontally oriented) and sinker (vertically oriented) roots were estimated from sap flow patterns. The vertical distribution of absorbing roots as derived from the analysis of sap flow patterns in stem sapwood was very similar to the distribution determined by the classical biometric analysis of fine roots. Trees leaning to the East had stem radii at the stump level and crown radii enhanced in the leaning direction. Sinker roots showed higher absorption activities in the leaning direction, but superficial roots were more absorbing in the opposite direction. The application of the above-described method allows for a better evaluation of the whole-tree behavior and facilitates the evaluation of tree and stand properties in traditional forest stands, which are not equipped for detailed scientific research. This may also facilitate practical applications in landscape-level studies.     

Reverse flow in roots of Sesbania rostrata measured using the constant power heat balance method

This investigation was performed to examine qualitatively and quantitatively the reverse flow in partially dried roots of Sesbania rostrata using the constant power heat balance method. First, a semi-empirical technique for estimating sheath conductance of sap-flow sensors without assuming that sap flow is zero at night was proposed. Sap flow measured with the heat balance method was compared with water uptake as measured by a potometric method. Sap flow was overestimated by 56·1% for a 3·3-mm-diameter root, and by 40·0% for 6·1 mm and 33·3% for 8·8 mm roots. However, high correlation coefficients between the rates of water uptake and sap flow demonstrated that calibration would provide reliable values for root sap flow. To detect reverse flow, a split root experiment was conducted using a S. rostrata plant with its root system divided between dry and wet compartments. Daily sap flow of the drying compartment declined whereas that in ‘wet’ root increased, suggesting that the decrease in water uptake by ‘dry’ roots was offset by the ‘wet’ roots. Reverse flow was observed at night in the root on the dry side of the container when the soil water potential was less than –0·30 MPa. The total amount of water released into the soil during the night period was estimated to be 22·5 g.     

华北山区核桃液流变化特征及对不同时间尺度参考作物蒸散量的响应

核桃是华北山区重要的经济树种,但该地区水资源紧缺,因此研究核桃液流耗水变化及其对气象因子的响应对于加强核桃水分管理、确定水分承载力具有指导意义。2008—2010年连续3个生长季节观测分析了华北山区核桃单株液流变化特征,研究了不同时间尺度(日尺度、月尺度)核桃液流与气象因子、参考作物蒸散量(ET0)的关系,并利用2009—2010年数据建立了拟合方程,在此基础上用拟合方程的估算值与2008年液流实测值进行了对比验证。结果表明:核桃液流具有明显的时间变化特征,最大液流量出现在5、6月份,2008—2010年生长季节(4—9月份)核桃液流量分别为893.23 L、854.88 L和841.77 L,日平均液流量分别为4.96 L、4.75 L和4.68 L。液流年际差异主要是由年降雨量引起。核桃液流与不同时间尺度下气象因子、ET0均有较好的相关性,且随着时间尺度的增加,二者间的相关性变大。利用2009—2010年液流实测值与气象因子、ET0建立的拟合方程,对2008年液流值进行了拟合,并与实测值进行了对比验证,发现不同时间尺度下液流拟合值与实测值均具有较好的一致性。在不能实现对液流进行连续观测的情况下,可以结合当地的气象资料、ET0等因子,利用建立的拟合方程对树木液流进行估算,弥补缺失的数据,具有较高的精确性,可用于指导当地核桃水分管理。     

Field and laboratory observations on the foraging and feeding techniques of blue tits (Parus caeruleus) and coal tits (P. ater) in relation to their habitats

I have tried to show (a) that results in the literature suggest that blue tits and coal tits survive better in their usual habitats of broad-leaved and conifer woods respectively; (b) that the higher mortality in the ‘wrong’ habitat may be attributed at least in part to an inability to acquire an adequate supply of food; (c) from my own and other people's field observations, that wild coal tits and blue tits use different feeding techniques; (d) from my own laboratory experiments, that each species is ‘best’ at the feeding techniques which it in general uses in the wild, and also that each species prefers to use the feeding techniques at which it is ‘best’; (e) that the feeding techniques of coal tits may be adapted to exploit the type of food supply found in conifers, while those of blue tits may be adapted for feeding in broad-leaved trees.     

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.     

Functional Traits and Water Transport Strategies in Lowland Tropical Rainforest Trees

Understanding how tropical rainforest trees may respond to the precipitation extremes predicted in future climate change scenarios is paramount for their conservation and management. Tree species clearly differ in drought susceptibility, suggesting that variable water transport strategies exist. Using a multi-disciplinary approach, we examined the hydraulic variability in trees in a lowland tropical rainforest in north-eastern Australia. We studied eight tree species representing broad plant functional groups (one palm and seven eudicot mature-phase, and early-successional trees). We characterised the species’ hydraulic system through maximum rates of volumetric sap flow and velocities using the heat ratio method, and measured rates of tree growth and several stem, vessel, and leaf traits. Sap flow measures exhibited limited variability across species, although early-successional species and palms had high mean sap velocities relative to most mature-phase species. Stem, vessel, and leaf traits were poor predictors of sap flow measures. However, these traits exhibited different associations in multivariate analysis, revealing gradients in some traits across species and alternative hydraulic strategies in others. Trait differences across and within tree functional groups reflect variation in water transport and drought resistance strategies. These varying strategies will help in our understanding of changing species distributions under predicted drought scenarios.     

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。     

马占相思夜间树干液流的分配及其对整树蒸腾估算的影响

采用Granier热消散探针测定了马占相思(Acacia mangium)的树干液流,结合Li-6400光合测定系统测定的夜间叶片气孔导度和蒸腾,将夜间液流区分为夜间树干水分补充和叶片气孔蒸腾。叶片的蒸腾作用微弱,因此,夜间液流主要用于补充贮水部位的水分亏缺。马占相思夜间水分补充量年内和年际的变化不明显,树形特征的差异是解释夜间水分补充量变化的重要因子,夜间水分补充量对于整树蒸腾量的贡献因季节和树木径级的不同而有明显变化,但对整树总蒸腾量计算造成的误差可以忽略。     

栓皮栎人工林树干液流对不同时间尺度气象因子及水面蒸发的响应

于2006-2008年主要生长季节,利用热扩散技术连续测算得到了华北土石山区30年生栓皮栎人工林液流量(SF),并结合同步测定的太阳辐射(Ra)、空气温度(Ta)、饱和水汽压亏缺(VPD)、风速(V)和降雨量(P)等气象因子、水面蒸发(EV0)及叶面积指数(LAI)等因子,分析了栓皮栎液流对不同时间尺度气象因子及水面蒸发的响应规律,探索建立长时间尺度水面蒸发与树干液流之间的关系模型。试验结果表明:(1)在主要生长季(4—9月份)期间,栓皮栎单株液流与同期测定的Ra、Ta、VPD和V等气象因子间均存在着较强的相关性。时间尺度为10min、1h、d、旬、月时,决定系数分别为0.388、0.482、0.539、0.654和0.812。说明随着时间尺度的增加,相关性越强;在不同时间尺度下,影响SF的最主要气象因子均为Ra。月SF与月Ra变化趋势的同步性尤为明显。(2)日尺度和月尺度上EV0与SF之间具有很好的线性关系,决定系数分别为0.578和0.876,比同时期、相等样本数条件下SF与Ra、Ta、VPD、V多元线性拟合的决定系数分别高3.6%和3.9%。(3)2006、2007、2008年生长季节降雨量分别为464.8、393.3mm和315.0mm,栓皮栎单株液流分别为2024.1L、1739.2L和1688.7L,年际间SF变化趋势与降雨量存在一定的一致性。     

基于热消散探针法的植物水分运输的研究进展

液流是分析树木耗水特性、研究树木水分传输机制的重要途径之一,热消散探针法已广泛用于树干液流变化的监测.热消散探针法是目前研究不同时空尺度上植物蒸腾耗水特性较为灵活、可靠、经济的一种方法.但由于物种特性的差异,可能造成试验过程中出现测量值与实际值相比偏低的状况.此外,相当一部分植物依赖树干储存水进行蒸腾,因此木材含水量的...     

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.     

Sap flow-based quantitative indication of progression of Dutch elm disease after inoculation with Ophiostoma novo-ulmi

Key message

The rate of progression of Dutch elm disease can be continuously and quantitatively estimated from sap flow measurements.

Abstract

Response of sap flow to inoculation with Ophiostoma novo-ulmi, a causal agent which causes vascular mycosis called Dutch elm disease, was studied in a field experiment comprised of 4-year-old wych elm trees (Ulmus glabra). Sap flow was measured on inoculated trees using the trunk heat balance method with external heating (EMS 62, Czech Republic) throughout the experiment. The first detectable symptoms of reduction in sap flow occurred 6 days after inoculation and all inoculated trees died within 16 days. Our experiment confirmed the ability of O. novo-ulmi to quickly kill young elm trees. The disease progressed faster than in previous experiments utilizing O. ulmi. To the best of our knowledge, this is the first experiment using sap flow measurements on trees inoculated by O. novo-ulmi. The trunk heat balance sap flow method is an effective non-invasive tool for continuous quantitative monitoring of the progression of vascular tree diseases, and show increased potential for field and greenhouse studies on changes in xylem hydraulic conductivity in a wide range of broadleaved and coniferous tree species.     

Spring Sap of Trees

In the spring, sap in perennial plants becomes available after winter dormancy. As is well known, substances including minerals and organic nutrients are transported from storage compartments in roots and stems to growth regions. For this solute distribution, xylem as well as phloem are used and probably also the cambial region and the ray system. In the present study, a puncture method has been used to record sap availability in six north European tree species during the spring. Attempts were made to withdraw sap from the trees at points 0.5 m, 1.3 m and 3.0 m above ground with “phloem needles” (constructed according to Hammel 1968), and a syringe. Sap could be withdrawn for the longest period for the sycamore (71 days). Birch (47 days) gave an intermediate period, beech (12 days), oak (12 days) and alder (13 days) gave short periods, and sap could not be withdrawn from ash. Effects of two environmental factors, temperature and light intensity on sap availability were examined. Temperature dependence of sap availability, already observed in the genus Acer, was confirmed in sycamore maple (Acer pseudoplatanus). There also seemed to be a correlation between sap availability and light intensity. Analysis of sugar in the tree sap revealed that sucrose was the only sugar in the sap of sycamore but hexoses (glucose and fructose) were preponderant in the sap of the other trees. The pH of sycamore sap (pH 6.9–5.4) fell as the season advanced. However, the pH-values of sap from the other trees did not vary significantly.     

Changes with seasonal flooding in sap flow of the tropical flood-tolerant tree species, <Emphasis Type="Italic">Campsiandra laurifolia</Emphasis>

In order to determine how flooding affects sap flow and hydraulic conductivity of the tolerant species, Campsiandra laurifolia, trees growing in a tropical seasonally flooded forest in Venezuela were studied. We hypothesized that trees respond to rising-waters with a decrease in root-water absorption, caused by hypoxia, and stomatal conductance, and that this is reverted later on through a process of acclimation that involves improvement in water absorption. We followed the seasonal changes, of trees with the whole or part of the canopy exposed to air, in sap flow density, leaf stomatal conductance, leaf transpiration rate and xylem water potential. The highest daytime sap flow density occurred at noon and its proportion relative to the yearly maximum (drainage at falling-waters) was 41 (dry season), 15 (flooding by rising-waters for 2 weeks), 54 (2 months of flooding) and 41% (6 months of flooding). Since at rising-waters dawn xylem water potential remained high, it became apparent that the initial stages of flooding imposed a restriction to sap flow unrelated to water deficit. The decrease at rising-waters in highest daytime sap flow density was due to reduced leaf-specific hydraulic conductivity, whereas the recovery observed 1.5 months later was correlated to an increase in hydraulic conductivity, and attributed to acclimation. Sap flow density was highly and positively correlated with radiation at all seasons but rising-waters; also, the relationship with air water vapor saturation deficit was high and significant on dates other than at rising-waters. Results suggest that early flooding inhibited water absorption by roots and that this inhibition was overcome later on at a higher water column through an acclimation process involving the improvement of internal aeration by adventitious roots.     

干旱区枸杞树干液流变化特征及其影响因素

树干液流作为植物蒸散作用的水分来源,是植物水分消耗的直观量化监测指标,利用包裹式树干液流监测技术获取干旱区枸杞全生育期树干液流实时数据,分析了不同时间尺度树干液流变化特征及各气象要素对树干液流的影响,为明晰枸杞耗水规律及其影响因素提供了重要的佐证。结果表明:枸杞的树干液流量昼夜差异较大,白天液流量是夜间的10倍左右;晴天液流速率、日累积量及变化幅度均大于阴雨天气,晴天液流速率变化曲线且呈宽峰型,在06:30左右启动较阴天提前30min;夏季树干液流启动时间为6:00比秋季提前1h左右,夏季的峰值123g/h。盛果期液流速率最大10.32g/h,营养生长期最小1.35 g/h;6—8月旺盛生长季,平均日耗水1388.3g/d,5—11月全生育期日均耗水1102.7g/d;树干液流速率与太阳辐射、空气温度均呈极显著正相关关系,与相对湿度呈负相关关系;枸杞树干液流(F)与太阳辐射(S)、温度(T)、相对湿度(H)及饱和水汽压(VPD)符合方程F=41.5+0.167S-0.563H+1.36T-9.67VPD(R~2=0.6547)。     

大连4种城市绿化乔木树种夜间液流活动特征

夜间液流有助于树木物质运输及其体内水分的补充(water recharge), 它不仅对植物的生长发育具有重要的生理生态学意义, 而且对大尺度植物蒸腾耗水的估算可能产生重要影响。2008年6月1日至8月31日, 以热扩散探针(thermal dissipation probe, TDP)技术对大连市劳动公园内的雪松(Cedrus deodara)、大叶榉(Zelkova schneideriana)、丝棉木(Euonymus bungeanus)和水杉(Metasequoia glyptostroboides) 4种乔木树种的不同径阶样木树干边材液流进行了测定, 并结合同步土壤水分与小气候观测结果分析了树木夜间(18:00至次日5:00)液流特征。实验结果表明, 树木普遍存在可感夜间液流, 夜间液流总量占观测期液流总量的比例在样木个体间呈现显著差异, 其变化范围为0.44%-75.96%; 观测期雨天夜间液流波动活跃, 显著高于晴天, 其单日夜间液流总量可持平, 甚至高于日间液流。相关分析表明: 水汽压亏缺(vapor pressure deficit, VPD)和风速的变化与夜间蒸腾显著相关, 它们能够较好地解释液流变化(R² > 0.6); 树木夜间液流主要用于夜间蒸腾和自身水分补充, 夜间液流现象主要发生在前半夜, 后半夜液流平稳且极接近0, 夜间液流量与相应的日间流量(R² = 0.356, p = 0.00)及胸径(R²_Spearman > 0.80)显著相关, 说明植物本身的结构和生理特点也是影响树木夜间液流的重要因子。单株样木夜间液流占全天总蒸腾量的比例低于14.4%, 如不考虑夜间液流的影响, 根据日间液流通过尺度扩展推算的森林生态系统年蒸腾量可能偏低。