The response of an Eastern Amazonian rain forest to drought stress: results and modelling analyses from a throughfall exclusion experiment

Warmer and drier climates over Eastern Amazonia have been predicted as a component of climate change during the next 50–100 years. It remains unclear what effect such changes will have on forest–atmosphere exchange of carbon dioxide (CO₂) and water, but the cumulative effect is anticipated to produce climatic feedback at both regional and global scales. To allow more detailed study of forest responses to soil drying, a simulated soil drought or 'throughfall exclusion' (TFE) experiment was established at a rain forest site in Eastern Amazonia, Brazil, for which time-series sap flow and soil moisture data were obtained. The experiment excluded 50% of the throughfall from the soil. Sap flow data from the forest plot experiencing normal rainfall showed no limitation of transpiration throughout the two monitored dry seasons. Conversely, data from the TFE showed large dry season declines in transpiration, with tree water use restricted to 20% of that in the control plot at the peak of both dry seasons. The results were examined to evaluate the paradigm that the restriction on transpiration in the dry season was caused by limitation of soil-to-root water transport, driven by low soil water potential and high soil-to-root hydraulic resistance. This paradigm, embedded in the soil–plant–atmosphere (SPA) model and driven using on-site measurements, provided a good explanation ( R ² > 0.69) of the magnitude and timing of changes in sap flow and soil moisture. This model-data correspondence represents a substantial improvement compared with other ecosystem models of drought stress tested in Amazonia. Inclusion of deeper rooting should lead to lower sensitivity to drought than the majority of existing models. Modelled annual GPP declined by 13–14% in response to the treatment, compared with estimated declines in transpiration of 30–40%.     

黄土高原苹果园蒸腾导度大气驱动规律比较

植物蒸腾导度是表征土壤-植物-大气连续体（SPAC）中植物-大气间水汽传导过程、反映植物水分调控能力的一类重要变量,常见有冠层导度（G_c）、冠层气孔导度（G_s）与叶片气孔导度（g_s）,明确三者在反映冠层蒸腾过程时的异同或关联性对于理解植物水分利用机制具有重要意义。本研究基于对黄土高原果园苹果树生长季内树干液流（J_s）及环境因子的连续观测,计算了G_c、G_s及脱耦联系数（Ω）等变量,并与短期连续观测的叶片气孔导度（g_s）比较,分析了G_c、G_s和g_s在反映冠层蒸腾特征方面的异同及其关系。结果表明,日变化过程中G_s、g_s呈\"单峰\"型曲线,而G_c则呈\"先增后减,午后抬升\"的\"双峰\"型曲线。g_s与G_s存在较紧密的线性关系（R²=0.80）,但与G_c的线性关系较弱（R²=0.02）。G_c、G_s均随大气水汽压亏缺（VPD）的变化呈现确定的规律,其中,上边界函数呈递减的对数函数关系,平均值则符合先增后减的Log-Normal函数关系（R²>0.95）,拐点对应的VPD值分别为1.33和1.16 kPa。在一日内,G_s对VPD变化的响应过程与g_s对VPDL （基于叶片温度计算的水汽压亏缺）变化的响应过程总体一致,其一致性高于G_c对VPD变化的响应。整个生长季（4-10月）中果树的Ω平均值为0.12,随着Ω递减,G_c与G_s的线性相关性愈趋紧密,其斜率呈递增趋势,G_c越来越趋近于G_s。研究结果表明,在北方地区,基于树干液流的监测能较准确的推导整株并估算林分的冠层蒸腾导度。与实测g_s的变化过程比较,G_s比G_c具有更高的一致性,G_s可以作为描述苹果树水分利用过程响应大气驱动的更为恰当的变量。     

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.     

Estimation of hydraulic conductance within field-grown apricot using sap flow measurements

Using the heat pulse and other techniques, the hydraulic architecture of apricot trees was mapped out. The flows (overall flow, flow across the four main branches) and forces (water potential differences between xylem and leaves) measured allowed us to quantify hydraulic conductance of branches and of the root/soil resistance. The experiment was carried out in a commercial orchard of 11-year-old apricot trees (Prunus armeniaca L., cv. Búlida, on Real Fino apricot rootstock) during 1 week (October 27–November 3, 1998). Three representative trees with a cylindrical trunk divided into four main branches of different sizes, orientation and local microclimate were chosen for the experiment. Sap flow was measured throughout the experimental period. Twelve sets of heat-pulse probes were used, one for each main branch. The diurnal course of the environmental conditions, the fraction of the area irradiated and leaf water relations were also considered in each main branch. The relationships between leaf water potential, xylem water potential and transpiration were established for different branches and also for the total plant. Using the slopes of these regressions, total plant conductance, the hydraulic conductance of the stem and root pathway, the hydraulic conductance of the canopy and the hydraulic conductance of each branch were estimated. Our findings show that the root conductance and the canopy hydraulic conductance are similar in magnitude. Leaf hydraulic conductance per leaf area unit was similar for each of the four branch orientations, indicating that, while the light microclimate has a dominant influence on transpiration, in this case it had little effect on the hydraulic properties of the canopy.     

The effect of fertilization on sap flux and canopy conductance in a Eucalyptus saligna experimental forest

Land devoted to plantation forestry (50 million ha) has been increasing worldwide and the genus Eucalyptus is a popular plantation species (14 million ha) for its rapid growth and ability to grow well on a wide range of sites. Fertilization is a common silvicultural tool to improve tree growth with potential effects on stand water use, but the relationship between wood growth and water use in response to fertilization remains poorly quantified. Our objectives in this study were to determine the extent, timing and longevity of fertilization effects on water use and wood growth in a non‐water limited Eucalyptus saligna experimental forest near Hilo, HI. We evaluated the short‐ and long‐term effects of fertilization on water use by measuring sap flux per unit sapwood area, canopy conductance, transpiration per unit leaf area and water‐use efficiency in control and fertilized stands. Short‐term effects were assessed by comparing sap flux before and after fertilizer application. Long‐term effects were assessed by comparing control plots and plots that had received nutrient additions for 5 years. For the short‐term response, total water use in fertilized plots increased from 265 to 487 mm yr^?1 during the 5 months following fertilization. The increase was driven by an increase in stand leaf area accompanied by an increase in sap flux per unit sapwood area. Sap flux per unit leaf area and canopy conductance did not differ during the 5 months following fertilizer additions. For the last 2 months of our short‐term measurements, fertilized trees used less water per unit carbon gain (361 compared with 751 kg H₂O kg C^?1 in control stands). Trees with 5 years of fertilization also used significantly more water than controls (401 vs. 302 mm yr^?1) because of greater leaf area in the fertilized stands. Sap flux per unit sapwood area, sap flux per unit leaf area, and canopy conductance did not differ between control and fertilized trees in the long‐term plots. In contrast to the short‐term response, the long‐term response of water use per unit wood growth was not significant. Overall, fertilization of E. saligna at our site increased stand water use by increasing leaf area. Fertilized trees grew more wood and used more water, but fertilization did not change wood growth per unit water use.     

麦田冠层气孔导度的分层研究

小麦灌浆期和乳熟期冠层各层叶片上、下表面的气孔导度之间呈正相关关系;冠层不同层的叶片气孔导度从早到傍晚有平行变化的趋势,数值上存在较大的差异,一般从冠层上到下递减。经分析,这主要与冠层叶片接受的光强自上而下递减有关,且这时所对应的叶片水势自冠层上到下递增的幅度大。测算结果表明,冠层气孔导度白天亦呈明显的日变化,灌浆期的值大于乳熟期的值。     

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.     

华北落叶松冠层平均气孔导度模拟及其对环境因子的响应

冠层气孔导度是生态水文学研究中的一个重要参数,研究其对环境因子的响应,能为建立机理性的森林蒸腾模型提供理论依据.本文利用热扩散探针,于2005年5-9月,测定了六盘山叠叠沟小流域华北落叶松人工林树干液流及其同步的环境因子,计算了林分冠层平均气孔导度(gc)并构建了Jarvis形式的冠层平均气孔导度模型,分析了gc对光合有效辐射(PAR)、空气水汽压亏缺(DVP)和土壤相对有效含水率(REW)的响应.结果表明:该模型能有效地计算gc的日变化特征,计算与观测的gc决定系数R2为0.76 (n=952).gc对各环境因子有不同响应关系,并表现为非线性特征.其中PAR是gc的驱动因子,当PAR＜0.35 mmol·m-2·s-1时驱动作用明显,大于此值则驱动作用变小;DVP是gc的限制因子,随着DVP的增加gc降低;REW=41%是gc对土壤水分响应的一个关键阈值,当REW＞41%时,土壤水分对gc的影响较小,当REW＜41%时,土壤水分则成为gc的限制因子.     

控水条件下侧柏冠层气孔导度对土壤水的响应

建立了不同控水条件下(无降水、一半降水、自然降水和二倍降水)的侧柏样地,于2016年8月-2017年8月监测了样地土壤含水量(SWC)、降水量、液流密度(Js)、叶面积指数(LAI)和水汽压亏缺(VPD)等因子,分析SWC对侧柏冠层气孔导度(gs)的影响.结果表明:一半、自然和二倍降水样地的SWC与降水量呈正相关,SW...     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.     

华南荷木林冠层气孔导度对水汽压亏缺的响应

冠层气孔导度(Gs)是量化气孔在冠层尺度水平上表现的参数,能够表征森林冠层表面水汽和能量交换的动态.本研究利用Granier树干液流测定系统,连续监测华南地区荷木林的树干液流,通过尺度转换和扩展获得冠层蒸腾速率,结合微气象观测值,以Pen-man-Monteith公式计算了Gs,并比较不同土壤水分条件下Gs对水汽压亏缺的响应.结果显示,Gs与气孔气体交换方法实测的叶片气孔导度(gs)日变化相似,单位转换数值大小与实测gs数量级一致.Gs对水汽压亏缺的响应在干季和湿季有明显差别:(1)在土壤水分充足的湿季(土壤含水量θ ＞33％),Gs对水汽压亏缺的响应更敏感(偏相关系数-0.316),而在干季(θ＜23％)则对光合有效辐射的响应更敏感(偏相关系数0.885).(2)荷木林冠层-大气脱耦联系数(Ω)在湿季接近l,干季则较湿季小,说明湿季叶片的界面层较厚,水汽压亏缺对Gs影响较小,而光合有效辐射是控制Gs的主要环境因子.     

Concurrent measurements of stem density, leaf and stem water potential, stomatal conductance and cavitation on a spaling of Thuja occidentalis L.

Abstract Concurrent estimates of stem density, leaf and stem water potential, stomatal conductance and ultrasonic acoustic emissions (cavitations) in an excised sapling of Thuja occidentalis L. were made. As the sapling dehydrated in air, the decline in leaf water potential to about - 2.0 MPa was followed by apparent rehydration of the foliage while the stem showed no sign of rehydration. The rate of acoustic emissions peaked prior to the onset of rehydration which coincided with virtual stomatal closure. There was a significant decline in stem density until maximum foliage rehydration level was reached. From this point, leaf water potential, stem water potential and stem density continued a relatively slow decline while acoustic emission rate and stomatal conductance remained low. Removal of the bark and majority of foliage from the sapling resulted in increased cavitation and more rapid deelines in leaf and stem water potential and stem density.     

鼎湖山南亚热带天然针阔叶混交林臭氧吸收特征

针阔叶混交林是我国南亚热带针叶林向地带性常绿阔叶林演替的中间林分类型,为我国南亚地区主要森林类型,发挥着重要的生态系统服务功能。基于树干液流技术和对臭氧浓度的连续监测,评价该森林类型的臭氧吸收特征和能力有着重要的环境生态学意义。对鼎湖山天然针阔叶混交林优势种马尾松(Pinus manssoniana)、锥栗(Castanopsis chinensis)、木荷(Schima superba)和华润楠(Machilus chinensis)在自然环境条件下的臭氧吸收能力进行了分析研究。结果表明:在日尺度上,4个优势树种的冠层气孔对臭氧导度(GO_3)和臭氧吸收通量(FO_3)均呈单峰型曲线,其最大值的时间在干季(10月至竖年3月)比湿季(4月至9月)滞后;季节尺度上,臭氧浓度在湿季达到最大值48.94 n L/L,湿季GO_3、FO_3和年臭氧吸收累积量(accumulative stomatal O_3flux,AFst)均显著高于干季(P 0.01),华润楠的臭氧吸收能力最强,在干季和湿季可分别达1.11 nmol m~(-2)s~(-1)和1.71nmol m~(-2)s~(-1)。随着水汽压亏缺(VPD)增大,优势种GO_3降低。光合有效辐射(PAR)超过1500 umol m~(-2)s~(-1)时,优势树种GO_3和FO_3呈下降趋势。针阔叶混交林的年臭氧吸收累积量超过了保护森林树木所采用的临界阈值,可认为鼎湖山针阔叶混交林受臭氧危害的潜在风险较高。     

Quercus ilex Transpiration as Affected by a Prolonged Drought Period

The effect of an extended drought (from 1992 to 1995) on water relations was assessed on evergreen oak (Quercus ilex L.) in a dehesa ecosystem (Seville, Southern Spain). Diurnal and seasonal transpiration patterns were analysed at leaf (porometry) and whole-tree level (sap flow), focusing on the relationship between tree transpiration rates (E_t) and potential evapotranspiration rates (PET). Daily maximum E_t varied over the year, becoming higher between May and August, and lower between November and April. Annual E_t (169 – 205 mm y^–1) accounted for less than 40 % of annual rainfall. The prolonged drought did not affect the water relations of the Q. ilex, mainly due to strong stomatal regulation avoiding the loss of water. Stomatal control was found in all seasons, although it was stronger in summer. This behaviour leads to low water consumption and low E_t/PET ratios throughout the year (0.05 to 0.27).     

Physiological Indicators of Plant Water Status of Irrigated and Non-irrigated Grapevines Grown in a Low Rainfall Area of Portugal

Water is a key resource in commercial wine production and both large excesses and deficits have undesirable effects upon the amount and quality of the wine produced. A balance between the water requirements of a fully developed canopy and the induced stress necessary for the commercial quality of the wine must be reached. Thus we need a physiological indicator that integrates both soil and climatic conditions to use as a management tool. An experimental field was established in the eastern part of the Demarcated Region of Douro – Portugal, to study the effect of water supply on the quality of the musts produced and we need a physiological indicator that relates to the water use and stress of the grapevines (Vitis vinifera L.) and to the later evaluation of the effect of irrigation practices upon the quality of the musts. We chose as indicators sap flow, leaf water potential at pre-dawn (0600 h), mid-morning (1000 h), solar noon (1400 h) and sunset (1900 h), stomatal conductance and leaf transpiration both measured at mid-morning and at solar noon, and related them to our experimental treatments that induce differences in soil water content, evaluated with time-domain reflectometry probes, with the objective of selecting the indicator that best describes the plant water status under different amounts of available water. Sap flow, leaf water potential and leaf transpiration rate measured at solar noon had highly significant correlations with soil water content and their regression on soil water content was also highly significant. Each of these parameters has shortcomings and none has a clear advantage over the other two as an integrator of the environmental conditions under these experimental conditions. Further studies of the parameters and their relationship with the quality characteristics of the produced musts are needed to achieve the ultimate objective of manipulating the soil water content.     

Water relations of baobab trees (Adansonia spp. L.) during the rainy season: does stem water buffer daily water deficits?

Baobab trees are often cited in the literature as water-storing trees, yet few studies have examined this assumption. We assessed the role of stored water in buffering daily water deficits in two species of baobabs (Adansonia rubrostipa Jum. and H. Perrier and Adansonia za Baill.) in a tropical dry forest in Madagascar. We found no lag in the daily onset of sap flow between the base and the crown of the tree. Some night-time sap flow occurred, but this was more consistent with a pattern of seasonal stem water replenishment than with diurnal usage. Intrinsic capacitance of both leaf and stem tissue (0.07-0.08 and 1.1-1.43 MPa(-1), respectively) was high, yet the amount of water that could be withdrawn before turgor loss was small because midday leaf and stem water potentials (WPs) were near the turgor-loss points. Stomatal conductance was high in the daytime but then declined rapidly, suggesting an embolism-avoidance strategy. Although the xylem of distal branches was relatively vulnerable to cavitation (P50: 1.1-1.7 MPa), tight stomatal control and minimum WPs near--1.0 MPa maintained native embolism levels at 30-65%. Stem morphology and anatomy restrict water movement between storage tissues and the conductive pathway, making stored-water usage more appropriate to longer-term water deficits than as a buffer against daily water deficits.