Granier树干液流测定系统在马占相思的水分利用研究中的应用

介绍了Granier热消散探针在树干液流测定中的工作原理,并利用该系统长期监测广东鹤山马占相思林14株样树的液流密度,分析了树木个体内和个体之间液流密度的差异、整树和林段水分利用的量化特征.由于树木边材结构以及周围微环境的差别,树木内和个体间的液流密度差异非常明显,变异系数的平均值分别为15.51%-37.26%、37.46%-50.73%.尽管液流密度的差异较大,但同一株树木不同方位的液流密度之间却呈现明显的线性相关（p＜0.0001）,这是重要的特征值,使得只需测定某一方位的液流密度经尺度外推计算整树和林段蒸腾成为可能.树木液流对环境因子响应的变化规律取决于所参照的时间尺度,日变化主要受光辐射、水汽压差等气候因子的控制,而土壤水份对液流的季节变化影响较大.形态特征明显影响树木的液流,高大树木由于边材较厚、树干粗壮和冠幅较宽而承载较多的辐射能量,因而水分蒸腾较高.对树木液流密度在径向和方位上进行适当的整合,可较准确地计算整树和林段蒸腾.由液流估测的马占相思整树和林段蒸腾的结果显示,该群落的水分利用在时间和空间上均有明显的分化.     

Influence of tree transpiration on mass water balance of mixed mountain forests of the West Carpathians

Brief information about water balance of the Carpathian temperate forest ecosystem are presented in the paper. Experimental research was done in a mature mixed fir-spruce-beech stand in the research plot “Pol’ana-Hukavský grúň” (850 m a.s.l.) in the south-eastern part of Pol’ana Mts. in the Biosphere Reserve UNESCO in Central Slovakia. Individual parameters of water budget have been continuously monitored. The water consumption of the model beech trees, as well as approximate model beech stand transpiration was estimated on the basis of sap flow measurements and up-scaling through dendrometrical approach. Sap flow of model beech trees was estimated by direct, non-destructive and continuous measurements by tree-trunk heat balance method with internal heating and sensing of temperature. These values were compared with potential evapotranspiration according to Türc. Precipitation parameters (rain and snow precipitation, through-fall precipitation, stem-flow, fog/snow precipitation and infiltration) have been measured simultaneously. Results of mass water balance and the portion of the tree transpiration within the individual water flows are presented. Evapotranspiration of beech-fir forest ecosystem in the middle mountain region (850 m a.s.l.) includes: transpiration (35% of precipitation total), interception (21%), evaporation (8%). There are differences between tree species in mass of transpirated water. Transpiration of spruce and fir reaches two-thirds of beech transpiration. Fog precipitation contribution to the water balance of beech-fir stand is 5%. Concurrently fog precipitation lowers the interception losses of vertical precipitation.     

油松、栓皮栎树干液流速率比较

应用TDP(ThermalDissipationProbe)技术对油松和栓皮栎树干液流进行了初步研究,经过野外近1a的实地定位观测,研究结果显示:栓皮栎月平均树干液流速率在整个生长期都较油松的月平均树干液流速率要高。前者大约是后者的5~10倍。栓皮栎在土壤干旱时期能够在白天产生明显的树干液流。在土壤干旱时期油松白天不产生树干液流而在晚上产生明显树干液流。在土壤相对湿润时期,油松和栓皮栎树干液流速率的波形与太阳总辐射的波形变化一致,但不同的是油松的树干液流速率波形呈明显的单峰状,而栓皮栎树干液流速率波形呈明显的多峰状。在土壤相对湿润时期太阳总辐射很低时能对油松树干液流速率产生明显的降低作用,而对栓皮栎树干液流则没有明显影响。在土壤干旱时期,油松和栓皮栎树干液流速率的峰值分别大约为0.0001cm/s和0.0006cm/s左右;在土壤水分充足时期,油松和栓皮栎树干液流速率的峰值分别大约相等约为0.0015cm/s左右,分别是油松和栓皮栎在干旱日期的液流速率峰值的10倍和2.5倍。     

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     

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.     

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。     

苹果树蒸腾规律及其与冠层微气象要素的关系

采用由热扩散植物液流技术测算得到的、周期为2 0 0 3年1～12月、时间步长10 min的苹果树蒸腾数据,结合同步观测得到果树冠层微气象要素值,分析太行山低山丘陵区10年生苹果树蒸腾耗水规律及其与微气象要素关系,旨在为该地区苹果生产提供必要的水分生态理论依据,并力图进一步完善苹果树蒸腾耗水理论。结果表明:(1)苹果树蒸腾(Tr)具有明显时间变化特征。Tr在晴天-多云日的主要和非主要生长季节日变化都表现为单峰曲线趋势。在阴天日,主要生长季节表现为多峰曲线趋势,非主要生长季节单峰除10月和3月份表现为双峰曲线趋势外,其余各月均为单峰曲线趋势。Tr日际变化或季节变化特征是3～4月份逐渐升高、5～6月份达到高峰值、7月份逐渐降低、10月末及11月初Tr迅速降低。全年Tr值为6 0 0 .9mm ,其中,主要生长期间(4～9月份)、非主要生长期间(1～3月份、10～12月份) TR值分别为5 0 2 .6、98.3mm ,分别占全年的83.6 %、16 .4 %。(2 )主要生长期内,Tr与冠层净辐射(Rn) ,空气温度(Ta)、湿度(RH) ,风速(V)等气象要素有很好的复相关性,统计回归方程为Tr =0 .2 74 0 .0 4 2 V 0 .0 0 6 7Ta - 0 .0 14 RH (n=2 6 35 2 ,Tr为单株蒸腾量,Tr、V、Ta、Rn的单位分别为L/ h、m/ s、℃、w/ m2 ,RH为无量纲值,以%表示) ,R2 =0 .     

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.     

Transpiration of trees and forest stands: short and long-term monitoring using sapflow methods

We show that sapflow is a useful tool for studies of water fluxes in forest ecosystems, because (i) it gives access to the spatial variability within a forest stand, (ii) it can be used even on steep slopes, and (iii) when combined with eddy correlation measurements over forests, it allows separation of individual tree transpiration from the total water loss of the stand. Moreover, sapflow techniques are quite easy to implement. Four sapflow techniques currently coexist, all based on heat diffusion in the xylem. We found a good agreement between three of these techniques. Most results presented here were obtained using the radial flow meter (Granier 1985). Tree sapflow is computed as sap flux density times sapwood area. To scale up from trees to a stand, measurements have to be made on a representative sample of trees. Thus, a number of trees in each circumference class is selected according to the fraction of sapwood they represent in the total sapwood area of the stand. The variability of sap flux density among trees is usually low (CV. 10–15%) in close stands of temperate coniferous or deciduous forests, but is much higher (35–50%) in a tropical rain forest. It also increases after thinning or during a dry spell. A set of 5–10 sapflow sensors usually provides an accurate estimate of stand transpiration. Transpiration measured on two dense spruce stands in the Vosges mountains (France) and one Scot's pine plantation in the Rhine valley (Germany) showed that maximum rate was related to stand LAI and to local climate. Preliminary results comparing the sapflow of a stand of Pinus banksiana to the transpiration of large branches, as part of the BOREAS programme in Saskachewan, Canada showed a similar trend. For modelling purposes, tree canopy conductance (g_c) was calculated from Penman-Monteith equation. In most experiments, calculated canopy conductance was dependent on global radiation (positive effect) and on vapour pressure deficit (negative effect) in the absence of other limiting factors. A comparison of the vapour pressure deficit response curves of g_c for several tree species and sites showed only small differences among spruce, oak and pine forests when including understorey. Tropical rainforests exhibited a similar behaviour.     

祁连山青海云杉低液流特征及其影响因素

植被蒸腾作为林区生态水文的重要组成部分,是分析林区水分循环及其植被健康生长的基础,对林区保护和科学管理至关重要。本研究于2017—2018年运用热扩散探针对祁连山青海云杉树干液流进行连续监测,探究祁连山青海云杉蒸腾耗水特征,并分析影响云杉生长和蒸腾过程的主控因素。结果表明: 青海云杉树干液流的瞬时变化在晴天呈单峰曲线,在阴天呈多峰或双峰型曲线,在雨天基本无明显规律。青海云杉液流密度与太阳辐射变化趋势一致,晴天液流启动较早,结束较晚,液流历时12～14 h。因该地区海拔较高(2700 m),空气温度较低,饱和水汽压差(VPD)较低,导致液流密度整体偏低,平均为(0.86±0.49) kg·d^-1。在小时尺度上,液流瞬时速率受太阳辐射和VPD的显著影响,而在日尺度下,0～40 cm土层土壤温度和土壤水分含量与液流密度显著相关。云杉液流密度随着太阳辐射、空气温度和VPD的降低而降低,在祁连山高海拔林区,较低的土壤和空气温度以及较低的VPD和太阳辐射是导致该地区青海云杉液流偏低的主要原因。     

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。     

北京山区元宝枫夜间液流活动特征及影响因素

树木夜间会维持部分气孔开放,从而能够在一定环境驱动因子的情况下进行夜间蒸腾。夜间液流作为储存水的重要来源,能够补充植物白天的水分亏缺,使其恢复水分储备,对植物生长发育有重要意义。采用TDP热探针法测定了位于八达岭林场的元宝枫树干液流密度,同步监测了主要环境因子,以深入揭示树木夜间蒸腾耗水规律和植被应对环境胁迫的调控机制,为山区植被建设、森林健康经营和挑选节水树种提供理论依据。结果表明:以0:00为界区分前半夜和后半夜,元宝枫夜间液流速率前半夜较后半夜活跃,且前半夜夜间累积液流量占夜间累积液流量的53.85%—64.10%,而后半夜夜间累积液流量占夜间累积液流量的35.9%—46.15%。5月的夜间累积液流量最大,平均夜间液流通量为5月6月8月9月7月。存在水分胁迫的条件下降雨之后夜间液流会增大,而当土壤水分条件较好,土壤水分不再是夜间液流的限制因子时,夜间液流通量并不高。不同树木形态的夜间液流通量有显著差异,在一定范围内,胸径树高冠幅越大的样木,夜间液流通量越大。用于夜间蒸腾的夜间液流通量与饱和水汽压差、温度、空气相对湿度、风速相关,其中夜间蒸腾存在于前半夜,表现为前半夜夜间液流通量与环境因子的相关性相较后半夜相关性较为显著,后半夜则以补水为主,补水量取决于土壤含水量和日蒸腾强度。存在干旱胁迫的条件下,夜间液流既用于夜间蒸腾,又有一部分用来补水;而土壤水分条件好时夜间液流则主要用于补水,此时夜间树干液流与环境因子相关性不高。元宝枫夜间液流通量的日蒸腾贡献率5、6月份大于7、8月份,即干季比湿季贡献率更高。夜间液流通量的日蒸腾贡献率与白天总蒸腾量相关性较高,并与累积太阳辐射成负相关。     

Tree water dynamics non-destructively assessed through sap flow measurements and potential evapotranspiration

Sap flow and potential evapotranspiration rates were analyzed for two coniferous tree species (Douglas-fir and Scots pine) and one broadleaf species (sessile oak) in a mixed Carpineto-Quercetum forest during the growing season 2005. The relationship between sap flow and potential evapotranspiration rates, effective crown area as a measure of the relative transpiration and daily relative proportion of the storage water used for transpiration were used as indicators of the tree water dynamics. These indicators were determined on four consecutive days and all three showed good reliability concerning tree water dynamics.     

Evaluation of the heat pulse velocity method for measuring sap flow in Pinus patula

Information on the water use of Pinus patula plantations isrequired to predict the impact of forest plantations on waterresources in South Africa. The heat pulse velocity (HPV) methodis a promising technique for measuring water use by trees, andhas been shown to measure sap flows accurately in a varietyof hardwood trees. This method has not been sufficiently verifiedfor pine trees where the presence of a strongly-defined ringstructure in the sapwood gives rise to a complex radial patternof sap flow. The purpose of this study was to compare wateruptake by cut trees to simultaneous HPV sap flow measurementsin the same tree. Fourteen trees were used for this comparison.Results showed that HPV sap flow estimates consistently overestimatedcut-tree uptake by an average of 49%. The bias is attributedto heat averaging across non-conducting latewood rings. Wateruptake was found to be highly correlated to the product of under-barkcross-sectional area and wound-corrected mean HPV, and it issuggested that this empirical relation provides a more appropriateway of estimating water use by this species. Key words: Heat pulse velocity, sap flow, Pinus patula, transpiration     

不同树高对荷木液流密度、整树蒸腾量和时滞的效应

人工林面积不断增大,这不仅能解决由于森林砍伐引起的一系列社会问题,而且还对解决水土保持、二氧化碳减排等环境问题起到重要作用。了解人工林的生长特性和蒸腾效率,对植被生长、恢复和管理有着重要意义。为此,该研究连续监测了华南地区12棵不同高度荷木人工林的液流密度,对样树以高度划分等级,采取错位相关法分析不同高度等级胸高处液流与冠层蒸腾的时滞效应。结果表明:气候环境相同时,所有样树胸高处液流日格型相似;荷木林蒸腾量优势木中间木劣势木,所有树木湿季月蒸腾量大于干季月蒸腾量;不同高度等级之间时滞差异显著,劣势木时滞50min,优势木和中间木时滞20min;所有样树干湿季时滞差异不显著,同一高度级两季节时滞差少于10min。这些说明:在干季华南地区土壤水分仍然相对较充足,植物输水阻力没有受到土壤水分降低和长距离水分传导的影响;中间木和优势木时滞短,水力阻力小,蒸腾量大并占据着林段的有利资源;劣势木长势低矮,时滞长,导管阻力大,蒸腾量少,光合作用需要的水热资源少,所以回馈根部的营养物质少,不均衡的营养循环使得林段分化愈明显,劣势木将逐渐从林段中被淘汰。该文指出在荷木人工林生长后期,对于长势低矮,生命力极弱的劣势木应定期砍伐,这样能增加优势木和中间木对光照及水分等有利资源的分配,提高林分质量,增加林地生产力。     

基于8hm2样地的天山云杉林蒸腾耗水从单株到林分的转换

为准确地估算天山北坡中部雪岭云杉森林的林分蒸腾耗水量,本研究设置了8hm2固定样地,在每木调查的基础上,通过对7株标准木连续晴天液流的观测,分别得到胸径与液流速率和边材面积的拟合方程,最终计算得到了林分的蒸腾耗水量,确定了估算该森林群落蒸腾耗水的最小取样面积,并且讨论了最小取样面积随样地位置的变化。结果表明：①在自然生长系,雪岭云杉林分最大蒸腾耗水量、平均蒸腾耗水量同植株密度呈Extrem函数分布;当种植密度达到1200株/hm2时,林分最大蒸腾耗水量达到最大,为77.27 t/hm2d;②对林分蒸腾耗水量的测算存在明显的最小取样面积,最小取样面积随取样点的不同而略有差异,取样起始点在本样地的海拔上边界和下边界处时最小取样面积为4 hm2,取样起始点在本样地的中部时最小取样面积为2.56hm2。本研究为更精确地估算天山北坡雪岭云杉森林的蒸腾耗水量提供了理论依据。     

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。     

Differences in transpiration of Norway spruce drought stressed trees and trees well supplied with water

The paper focuses on the evaluation of transpiration as a physiological process, which is very sensitive to drought stress. Reactions of 25-year-old Norway spruce (Picea abies (L.) Karst.) trees to drought were examined during 2009 summer. Sap flow rate (SF), meteorological and soil characteristics were measured continually. Vapour pressure deficit of the air (VPD) and cumulative transpiration deficit (KTD) was calculated. During the second half of the vegetation period, the decrease in soil water content was observed and irrigation was applied to a group of spruce trees, while the second group was treated under natural soil drought. On the days, when the differences in transpiration between irrigated (IR) and non-irrigated (NIR) trees were significant (21 days), transpiration of NIR trees was only 23% of the transpiration of IR trees. We found significant differences in transpiration when the soil water content (SWC) of NIR variant at a depth of 5–15 cm ranged from 10.4 to 13.7%. Under both regimes of water availability, daily transpiration significantly responded to atmospheric conditions. However, the influence of all assessed meteorological parameters on SF of NIR trees was significantly lower than on IR tree. The dependency of transpiration on evaporative demands of atmosphere decreased with the decreasing soil moisture. Cumulative transpiration deficit of the stand during the entire evaluated period was 50.9 mm. The difference between the transpiration of the mean NIR tree and of the mean IR tree was 278.8 L over the assessed period of 47 days (5.9 L per day). The transpiration of NIR trees was 40.3% from the transpiration of IR trees during this period.     

Spatial patterns of ground vegetation, soil microbial biomass and activity in a mixed spruce-birch stand

Trees directly and indirectly influence the above- and below-ground environment, and can be expected to modify the spatial patterns of organisms associated with the forest floor. This study aimed to examine the effects of a coniferous (Picea abies) and a broad-leaved (Betula pubescens) tree species on the spatial pattern of ground vegetation and soil microbial properties in a mixed stand in central Sweden. I have characterised the species composition of ground vegetation, soil microbial biomass and activity, photosynthetic active radiation (PAR), soil water content and soil pH in the stand, and tested whether the spatial patterns of these variables were related to the positioning of trees. Geostatistics were used to describe the spatial variation in ground vegetation, soil mirobiological properties and the soil surface environment. PAR, soil water content and the cover of the moss Brachytecium reflexum and associated herb species decreased with the influence of spruce trees. Microbial biomass, measured as the amount of phospholipid fatty acids, decreased with spruce influence but increased with the influence of birch trees. Microbial respiration was not affected by spruce but increased with the influence of birch. Ground vegetation and microbial respiration, which were influenced by one tree species only, aggregate on a scale of 4-5 m, corresponding fairly well with patches of a single tree species. Soil microbial biomass, which was affected by both tree species, aggregated on a scale of 7-8 m. roughly corresponding to the distance between patches of spruce and birch trees respectively. I suggest that spruce trees influenced vegetation mainly through shading, and that a difference in the availability of organic matter under birch and spruce trees caused spatial variation in microbial biomass and activity. Thus, spatial patterns in ground vegetation and soil microbial properties may develop in a mixed forest of coniferous-broad leaved trees, as a result of the difference in influence of tree species and nested variation associated with the arrangement of the trees.     

Branch transpiration of pine and spruce scaled to tree and canopy using needle biomass distributions

Branch water exchange and total tree water uptake were measured in a mixed Norway spruce and Scots pine stand in central Sweden during the 1995 and 1996 growing seasons. Branch transpiration was scaled to canopy level on the basis of a branch conductance model, using vertical needle-area distributions obtained by destructive sampling. Comparison with total tree water uptake scaled to canopy level showed agreement within 10%, for periods when the canopy was not affected by climatically induced stress. Comparison of scaled fluxes on individual trees showed that measurements of transpiration at branch level provide information on the direct response of transpiration to variations in weather, and furthermore that the time-lag between transpiration and tree water uptake was as much as 3 h. The vertical needle-area distribution of Scots pine was similar to that found by other authors. Needle-area distribution on Norway spruce, which has not been described before, showed that it has its largest needle area at the top of the crown. Specific needle area varied considerably both within trees and between trees. For spruce, mean specific needle area (±SD) varied from 2.4±0.5 mm² mg^–1 at the top of the crown to 7.1±1.9 mm² mg^–1 at the base. Corresponding figures for Scots pine were 3.4±2.0 and 9.1±2.1. Received: 5 March 1999 / Accepted: 17 March 2000