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.     

基于树干液流测定值的马占相思(Acacia mangium)冠层气孔导度计算及数值模拟

为深入揭示华南地区马占相思冠层气孔导度对环境因子的响应规律,在2005年7月至11月,利用Granier热消散式探针法对马占相思(Acacia mangium)的树干液流(sapflow)进行了连续测定,计算出整树的蒸腾,并由Penman-Monteith方程得出马占相思的冠层气孔导度值。通过分析,发现:马占相思冠层气孔导度是控制马占相思树整树蒸腾的主要因素;冠层气孔导度随着水汽压亏缺增加呈负指数函数下降的趋势。使用包括了太阳总辐射、水汽压亏缺和气温的Jarvis模型可以较好地模拟马占相思冠层气孔导度对环境因子的响应特征;模拟结果表明:环境变量对模型精确度的影响程度依次为:水汽压亏缺>太阳总辐射>气温。     

基于树干液流测定值的马占相思（Acacia mangium）冠层气孔导度计算及数值模拟

为深入揭示华南地区马占相思冠层气孔导度对环境因子的响应规律,在2005年7月至11月,利用Granier热消散式探针法对马占相思（Acacia mangium）的树干液流（sap flow）进行了连续测定,计算出整树的蒸腾,并由Penman-Monteith方程得出马占相思的冠层气孔导度值。通过分析,发现：马占相思冠层气孔导度是控制马占相思树整树蒸腾的主要因素;冠层气孔导度随着水汽压亏缺增加呈负指数函数下降的趋势。使用包括了太阳总辐射、水汽压亏缺和气温的Jarvis模型可以较好地模拟马占相思冠层气孔导度对环境因子的响应特征;模拟结果表明：环境变量对模型精确度的影响程度依次为：水汽压亏缺>太阳总辐射>气温。     

荷木整树蒸腾对干湿季土壤水分的水力响应

降雨在时间上的非均匀分配导致森林土壤含水量呈现明显的干、湿季变化,并可能在干季形成水分胁迫,引起植物蒸腾变化。在监测环境因子的同时,利用Granier热消散探针连续监测荷木(Schima superba)的树干液流,以液流密度值计算整树蒸腾,并结合水力导度与叶片/土壤的水势差,探讨环境因子和水力导度对荷木整树蒸腾的协同控制。结果表明,华南地区的季节性降雨形成的干、湿季并未引起荷木蒸腾在季节上的显著差异,但对产生蒸腾的水力生理产生了显著影响。荷木蒸腾在干、湿季均与主要驱动环境因子(光合有效辐射PAR和水汽压亏缺VPD)呈显著正相关。在水热充足的湿季,荷木蒸腾主要受气孔导度调节;在干季,当空气水汽压亏缺达2.132 MPa时,水力导度与气孔导度协同控制蒸腾。整树水力导度对整树蒸腾的水力补偿出现在15:00—17:00,平均补偿值为0.08 g/s。利用蒸腾的估测值与实测值之间的差值量化荷木的水力补偿效应,是对水力导度与气孔导度协同控制树木蒸腾机理的深入探索。研究结果对于掌握季节性降雨不均背景下华南地区主要造林树种需水和耗水规律,有效发挥森林保水功能具有重要意义。     

华北山区侧柏冠层气孔导度特征及其对环境因子的响应

冠层气孔导度(g_s)是衡量冠层-大气界面水汽通量的重要生物学常数,研究其特征及对环境因子的响应,能为开展森林冠层水汽交换过程的机理性研究提供理论依据.于2014年利用SF-L热扩散式探针测定了侧柏的树干液流密度(J_s),同步监测光合有效辐射(PAR)、饱和水汽压差(VPD)、气温(T)等环境因子,计算侧柏的冠层气孔导度特征并分析其对各环境因子的响应.结果表明: 侧柏液流密度的日变化总体呈双峰曲线,生长季高于非生长季,且胸径越大液流密度越大;冠层气孔导度日变化与单位叶面积冠层蒸腾(E_L)趋势相近,均呈双峰曲线,生长季的冠层气孔导度和蒸腾较非生长季略高.侧柏冠层气孔导度与空气温度呈抛物线关系,在10 ℃左右冠层气孔导度达到峰谷;光合有效辐射以400 μmol·m^-2·s^-1为界,小于该阈值两者呈正相关关系,大于该阈值则冠层气孔导度受其影响较小;与饱和水汽压差呈负对数函数关系,随饱和水汽压差增大而逐渐降低.较高的空气温度和光合有效辐射、较低的饱和水汽压差有利于侧柏形成较大的冠层气孔导度,进而促进冠层蒸腾.     

马占相思林冠层气孔导度对环境驱动因子的响应

利用Granier热消散探针在2003年10月测定了广东鹤山丘陵地马占相思林14株样树的树干液流,同时监测林冠上方的光合有效辐射、空气湿度和气温,结合树木的形态和林分的结构特征,计算马占相思的整树蒸腾(E)、林分总蒸腾(E_t)以及冠层平均气孔导度(g_c),分析树形特征与整树水分利用的关系、冠层气孔导度对光合有效辐射(PAR)和空气水汽压亏缺(D)的响应.结果表明,整树蒸腾与胸径(P＜0.0001)、边材面积(P＜0.0001)和冠幅(P=0.0007)以自然对数的形式、与树高(P=0.014)以幂函数的形式呈现显著正相关.冠层气孔导度最大值(g_cmax)随D的上升呈对数函数下降(P＜0.0001),对光合有效辐射的响应则呈双曲线函数增加(P＜0.0001).液流测定系统能提供连续和准确的整树和林分蒸腾速率值,经严格数学推导公式计算,最终可求出冠层气孔导度,是研究森林水分利用与环境因子相互关系的有效方法.     

宁夏河东沙区侧柏冠层气孔导度对环境因子的响应及其模拟

为了解林木冠层气孔导度对环境因子的响应规律,进一步阐释冠层气孔导度调控森林蒸腾的机理,本研究以宁夏河东沙区典型防护林树种侧柏(Platycladus orientalis)为对象,利用Granier热扩散技术,在生长旺季(7—9月)连续监测树干液流,并同步观测小气候;利用Penman-Monteith方程计算获得连续的冠层气孔导度(gc),并采用Jarvis-Stewart气孔模型对侧柏冠层气孔导度变化过程进行模拟。结果表明:环境因子与gc呈极显著相关关系(P0.01),对侧柏gc的影响从大到小依次为大气温度(Ta)太阳辐射(Rs)饱和水气压亏缺(VPD)空气湿度(RH)。典型晴天下,gc与VPD、Ta的关系曲线上升阶段和下降阶段路径不同,呈现"磁滞回环"的特性,且不同时段表现出不同的响应规律。Jarvis-Stewart模型较好地模拟了侧柏冠层气孔导度对环境变量Rs、VPD、Ta的响应过程,采用交叉验证法对模型进行验证和参数率定,可以解释84.4%的变量。     

北京山区侧柏林冠层-大气蒸腾导度模拟及环境因子响应

蒸腾导度模型是衡量冠层-大气界面水汽输出的重要阻力模型,研究其特征及对环境因子的响应,为揭示森林冠层-大气界面水汽输出阻力机制提供理论依据。以首都圈森林生态系统定位观测研究站侧柏林为研究对象,采用TDP热探针法测定侧柏林树干液流密度,同步监测光合有效辐射、饱和水汽压差、气温、风速等主要环境因子,分析冠层导度和空气动力学导度的动态变化,构建冠层-大气蒸腾导度模型并模拟,明确冠层-大气蒸腾导度对各环境因子的响应关系。结果表明:蒸腾导度季节变化表现为非生长季与冠层导度趋势一致,生长季与空气动力学导度趋势一致,全年均为单峰趋势。冬季蒸腾导度与冠层导度保持较稳定差值(45 mol m^(-2 )s^-1左右),其他季节蒸腾导度与冠层导度、空气动力学导度的最大差值,均在各季节冠层导度、空气动力学导度的峰值水平。全年日均蒸腾导度冬季最大(86.92 mol m^(-2 )s^-1),其他季节较小且稳定(40—50 mol m^(-2 )s^-1之间)。在非生长季各环境因子对蒸腾导度的影响与对冠层导度的影响基本一致,温度为主要影响因子(r=-0.198),其他环境因子影响较小(r<0.1);在生长季中风速为主要影响因子(r=0.488),光合有效辐射(r=0.228)和饱和水汽压差(r=-0.299)的影响明显升高,温度的影响降低(r=0.114)。蒸腾导度模型较好的模拟了冠层-大气界面侧柏蒸腾不同季节的变化规律,阐明了各环境因子和冠层导度、空气动力学导度对蒸腾导度的影响机制,证实在生长季应重视空气动力学导度对蒸腾的影响。     

树高对马占相思整树水分利用的效应

利用Granier热消散探针,于2004年观测了华南丘陵坡地常见绿化先锋树种马占相思(22年生)的树干液流,同时监测林冠上方的光合有效辐射、气温、相对湿度和0～30 cm的土壤体积含水量.结合树木的形态特征、液流密度和简化的Whitehead & Jarvis公式,分别计算了整树蒸腾、冠层气孔导度和叶面积/边材面积比值,分析了树高对整树蒸腾、冠层气孔导度和叶面积/边材面积比值的影响.结果表明：土壤水分充足时,马占相思整树蒸腾随树高呈二次多项式增加(P<0.01),冠层气孔导度日变化均呈“单峰”格型;在所有光合有效辐射范围内,高树的参比冠层气孔导度和冠层气孔导度对水汽压亏缺的敏感性均高于矮树;叶面积/边材面积比值为（1.837±0.048） m²·cm^-2,并与树高呈幂函数关系.随着树木高度的增加,马占相思没有发生明显的水力限制和补偿.     

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.     

基于树干液流测定值进行尺度扩展的马占相思林段蒸腾和冠层气孔导度

应用Granier热消散探针,长期监测华南丘陵地马占相思(Acacia mangium)林14棵样树的树干液流(Sap flow),由此计算整树和林段的蒸腾速率,结合同步记录的环境因子,求算冠层平均气孔导度(G_c)。Granier探针的灵敏度较高,能精确测定即使是微弱的液流活动。观测结果显示,树木个体之间的液流密度(J_s)和整树蒸腾(E_t)受树形特征影响较大。马占相思林径级大的树木个体数较少,但占据林段边材总面积和林段蒸腾的比例较大。J_s和E_t的日变化主要受光合有效辐射(Q_o)和空气水蒸气压亏缺(D)的控制,土壤含水量(θ)对较大胸径树木E_t的影响大于胸径较小的树木,个体之间J_s和E_t的差异随θ的下降而缩小。一年中,林段蒸腾(E)在光照和水热条件较好的7月最高,9~12月,由于土壤水分供应的减少致使E值降低,E对D的敏感性下降。G_c与主要环境因子的关系与E相似,如果θ长期偏低,G_c会明显下降,是造成E降低的主要原因。成熟马占相思林在光照充足、水热条件较好的情况下的蒸腾活动旺盛,但对土壤水分胁迫的忍受力较低。     

Coordination of stomatal,hydraulic, and canopy boundary layer properties: Do stomata balance conductances by measuring transpiration?

A striking coordination is observed in sugarcane between prevailing levels of stomatal opening and the hydraulic capacity of the soil, roots and stem to supply the leaves with water. This coordination of vapor phase and liquid phase conductances is associated with decreases in stomatal conductance on a leaf area basis that compensate for increasing leaf area during canopy development, causing transpiration to approach a maximum value on a per plant or ground area basis rather than increase linearly with leaf area. The resulting balance between water loss and water transport capacity maintains leaf water status remarkably constant over a wide range of plant. sizes and growing conditions. These changes in stomatal conductance during development are determined by changes in the composition of the xylem sap rather than by changes in leaf properties. Changes in boundary layer conductance resulting from non-developmental changes in canopy structure such as loding cause additional changes in stomatal conductance mediated by altered humidity at the leaf surface. These maintain a constant level of total canopy vapor phase conductance (stomatal and boundary layer in series) and a constant level of canopy transpiration. These patterns indicate that stomata exert an active role in regulating transpiration even in dense canopies. This control function is consistent with stomatal metering of transpiration, mediated by fluxes of root-derived materials in the xylem sap.     

北京山区油松和元宝槭冠层气孔导度特征及其环境响应

叶片气孔是植物进行水汽交换的通道, 影响着植物的蒸腾和光合作用。然而叶片气孔行为受环境条件和树种类型的影响, 不同树种冠层气孔导度对环境因子响应的差异性, 以及在生长季不同时期叶片气孔对冠层蒸腾的调节作用是否会发生改变, 仍不清楚。该研究目的是通过探究各环境因子对不同树种冠层气孔导度的相对贡献率以及叶片气孔对冠层蒸腾的调节作用, 为深入了解植物水分利用状况和山区森林经营提供参考依据。于2018年生长季以北京八达岭国家森林公园内的58年生油松(Pinus tabuliformis)和39年生元宝槭(Acer truncatum)为研究对象, 利用热扩散技术对其树干液流进行连续监测, 并同步监测环境因子。利用彭曼公式计算冠层气孔导度(G_s)。主要结果: (1)油松和元宝槭日间G_s在日、月时间尺度上存在明显差异。5-7月油松和元宝槭日动态G_s均随饱和水汽压差(VPD)和太阳辐射(GR)的增加呈上升趋势, 上升持续时间比8月和9月长; 在月尺度上, 随着VPD、GR的降低和土壤湿度(VWC)的升高, G_s从5月到9月整体上升。(2)利用增强回归树法分析得到VWC和VPD对G_s的贡献率最大, 其次是GR、气温和风速。VWC和VPD对油松G_s的贡献率分别为66.4%和17.4%, 对元宝槭G_s的贡献率分别为54.8%和21.0%。(3)油松和元宝槭的dG_s/dlnVPD值与参考冠层气孔导度之间的斜率均显著高于0.6, 气孔调节作用相对较强。综上所述, 气孔对环境因子的响应在树种以及生长季不同时期之间存在差异, 为防止水分过度散失, 两树种在不同土壤水分条件下均通过严格的气孔调节控制蒸腾量。     

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements

Summary Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day^–1 tree^–1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m² of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s^–1 that is equivalent to 112 mmol m^–2 s^–1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m^–2 s^–1 at the top and 150 mmol m^–2 s^–1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was –27.76±0.27 (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.This paper is dedicated to Prof. Dr. O.L. Lange on the occasion of his 65th birthday     

Correlations of stomatal conductance with hydraulic and chemical factors in several deciduous tree species in a natural habitat

Recent research in whole-plant stomatal physiology, conducted largely with potted plants in controlled environments, suggests that stomatal conductance ( g _s) might be more closely linked to plant chemical variables than to hydraulic variables. To test this in a field situation, seasonal g _s was examined in relation to a number of plant and environmental variables in 11 temperate, deciduous forest tree species. Stomatal conductance was generally better correlated with environmental variables (air temperature, vapor pressure deficit, PPFD) than with plant variables, and slightly better correlated with plant hydraulic variables (shoot water and osmotic potentials) than with plant chemical variables (xylem sap ABA concentration, xylem sap pH). We examined a model, developed previously for maize, which describes regulation of g _s by xylem sap ABA concentration with leaf water status acting to modify stomatal sensitivity to the ABA signal. This model explained slightly more variation in seasonal g _s in the forest trees than did single plant variables but not more variation than most single environmental variables. Response surface models, especially those incorporating environmental variables, were more consistently successful at explaining g _s across species.     

Stomatal control of transpiration in the canopy of a clonal Eucalyptus grandis plantation

 Predawn leaf water potential, stomatal conductance and microclimatic variables were measured on 13 sampling days from November 1995 through August 1996 to determine how environmental and physiological factors affect water use at the canopy scale in a plantation of mature clonal Eucalyptus grandis Hill ex-Maiden hybrids in the State of Espirito Santo, Brazil. The simple ”big leaf” Penman-Monteith model was used to estimate canopy transpiration. During the study period the predawn leaf water potential varied from –0.4 to –1.3 MPa, with the minimum values observed in the winter months (June and August 1996), while the average estimated values for canopy conductance and canopy transpiration fell from 17.3 to 5.8 mm s^–1 and from 0.54 to 0.18 mm h^–1, respectively. On the basis of all measurements, the average value of the decoupling coefficient was 0.25. During continuous soil water shortage a proportional reduction was observed in predawn leaf water potential and in daily maximum values of stomatal conductance, canopy transpiration and decoupling coefficient. The results showed that water vapour exchange in this canopy is strongly dominated by the regional vapour pressure deficit and that canopy transpiration is controlled mainly by stomatal conductance. On a seasonal basis, stomatal conductance and canopy transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture and rainfall. Good results were obtained with a multiplicative empirical model that uses values of photosynthetically active radiation, vapour pressure deficit and predawn leaf water potential to estimate stomatal conductance. Received: 10 June 1998 / Accepted: 20 July 1998     

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.