3类典型温带山地森林的叶面积指数的季节动态: 多种监测方法比较

 叶面积指数(leaf area index, LAI)是定量描述冠层结构的最有效指标之一。鉴于森林冠层三维结构的高度复杂性和异质性, 迄今仍没有形成统一标准的LAI测量方法。该文利用LAI-2000冠层分析仪、CI-110冠层分析仪和半球摄影法(digital hemispherical photograph, DHP), 对北京东灵山地区以蒙古栎(Quercus mongolica)为主的落叶阔叶林、华北落叶松(Larix gmelinii var. principis-rupprechtii)林和油松(Pinus tabuliformis)林的有效叶面积指数(effective leaf area index, LAI_e)进行了动态监测, 探寻其季节变化规律。为准确地估算温带山地主要森林类型的LAI, 对光学仪器测量值进行了去除木质成分、聚集效应等校正, 与基于凋落物收集法的相应实测值进行了比较分析。结果表明: 3种典型森林在生长季期间叶片生长均呈现单峰型; 3种光学仪器测量方法的同期LAI_e数值大小顺序为: LAI-2000冠层分析仪>DHP>CI-110冠层分析仪。光学仪器的直接测量值LAI_e包含了木质成分的贡献, 钝化了季节动态的变化幅度, 这对有明显季节交替的落叶林尤为突出。经校正, LAI-2000冠层分析仪和DHP的测量值与实测值都表现出显著的相关性, 其中LAI-2000冠层分析仪最适于采用基于空隙大小的校正方法, 而基于空隙度和空隙大小的综合算法则是校正DHP的最佳选择。结合经济成本和野外实际操作等因素考虑, DHP具有更大的推广优势, 特别适用于温带山地落叶林。     

叶面积指数田间测量中有限长度平均法的改进

叶面积指数(LAI)的田间测量是生态和农业等领域的常规工作之一, 测量方法分为直接测量和间接测量, 间接测量中有一类方法基于数字相机照片提取冠层孔隙率, 再用有限长度平均法同时估算LAI和聚集指数。然而, 有限长度平均法自提出以来缺少进一步的发展, 在有限长度的样线/样方上应用比尔定律的方式具有理论缺陷, 可能造成无效值或高估LAI。从模拟的训练数据中提取经验公式以取代比尔定律进行样线/样方的LAI估算, 提高了有限长度平均法的精度和鲁棒性。进一步分析在一定精度需求下对样线/样方大小和数量的要求, 对于非均匀样地, 提出样线长度为8倍等效叶片边长、样方边长为3倍等效叶片边长的推荐设置。在基于数字相机照片提取非均匀样地LAI的应用中, 使用样方采样比样线采样更为适宜。     

落叶收集法测定叶面积指数的快速取样方法

叶面积指数(LAI)是植被冠层结构的一个重要参数,它不仅是许多生态和气候模型的重要输入变量,而且是生态系统动态变化监测的一个重要指标。LAI可通过各种间接和直接的手段来观测,而间接观测的LAI值常常需要直接的观测数据来校验。落叶收集法是一种广泛使用的直接观测LAI的方法,过去的研究还未发现有涉及落叶收集的取样技术及其观测精度的内容。对长白山和北京地区落叶阔叶林的落叶进行了3a的观测,每年一次性收集落叶样品分析,研究结果表明:①不同层次落叶的含水量差异巨大,且落叶含水量的日变化明显。上下层落叶的含水量绝对值差异高达10%以上,日变化绝对值差异高达20%以上。因此,在野外收集落叶样本时,为减小落叶含水量变化所导致的LAI观测误差,应从上到下直到地面进行取样,且尽可能多地收集落叶样本。②在落叶阔叶人工林和天然林里,无论样地的大小(1hm2或30m×30m样地),无论取样单元的大小(1m2或25m2分辨率),林内的LAI分布很不均匀,LAI介于0-15.5(1m2分辨率的1hm2样地)或者2.6-9.1(25m2分辨率的30m×30m样地)。③要准确测定落叶林的LAI,收集落叶的样地面积越大越好,且尽量选择地势平坦的样地。对于1hm2或者30m×30m大小的样地,可随机布设一个10m×10m的小样地来观测,精度分别可达85%、80%。④10m×10m小样地的LAI观测,可将其分为4个相邻的5m×5m小样进行取样。对每个5m×5m小样,快速的取样方法是:Ⅰ.随机布设6个1m2取样,这样取样可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为90%、75%、70%左右。Ⅱ.随机布设11个1m2取样。可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为94%、80%、75%左右。     

南方人工林叶面积指数的摄影测量

利用CI-110冠层分析仪、鱼眼镜头数码照片分析,结合胸径-叶片半表面积模型对千烟洲不同森林类型(20年生)的叶面积指数进行了测定,并对3种方法进行了比较.共调查74块样地.研究对象主要为湿地松(Pinus elliottii)林、马尾松(Pinus massoniana)林、杉木(Cunninghamia lanceolata)林.针对上述3种林型,CI-110观测值分别为1.355(0.38～2.85)、1.265(0.62～2.55)和2.140(0.76～3.31);鱼眼镜头测定值依次为1.339(0.71～1.79)、1.491(1.11～1.92)和1.984(1.26～2.59).利用胸径-叶片半表面积模型测定的叶面积指数(范围)依次为4.53(2.24～6.34)、3.88(1.19～6.28)和6.63(3.5～8.37).间接仪器法测定的值明显小于实测值,但具有一定的相关性.胸径-叶片半表面积模型测定叶面积指数最为可靠,而鱼眼镜头测定法比冠层分析仪测定精度高.     

Estimation of deciduous forest leaf area index using direct and indirect methods

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the Demon portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI–WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.     

祁连山区青海云杉林冠层叶面积指数的反演方法

叶面积指数(Leaf area index, LAI)是陆地生态系统的一个十分重要的结构参数。随着空间精细化模型的发展和基于过程的分布式模拟技术的应用, 对LAI的区域估算显得越来越重要, 但目前尚缺乏有效的估算手段。该项研究以青海云杉(Picea crassifolia)林为研究对象, 利用LAI-2000冠层分析仪、鱼眼镜头法和经验公式法对林冠层LAI进行了测定, 观测值分别为1.03~3.70、0.48~2.26和2.27~8.20, 显然, 仪器测定值偏低。针对针叶的集聚效应导致仪器测定值偏低的现象, 利用跟踪辐射与冠层结构测量仪(TRAC)测定的青海云杉林聚集系数计算调整系数, 对鱼眼镜头法获取的LAI值进行订正。根据高分辨率的遥感数据反演青海云杉林的植被指数与LAI的关系, 最后获得了较合理的该地区林冠层LAI的空间分布图。     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

云冷杉针阔混交林叶面积指数的空间异质性

叶面积指数(Leaf area index,LAI)是森林生态过程的关键参数和描述森林冠层结构的重要指标。用半球摄影技术对吉林省汪清林业局金沟岭林场的10块1 hm2云冷杉针阔混交林的LAI进行测定,采用地统计学的半变异函数和普通克里格法对研究区的LAI的空间异质性进行了分析。结果表明:10块样地的10 m×10 m小样方内以及样地间的LAI离散程度较小,但分布有一定的规律。LAI的空间相关性存在不确定性,可由线性、孔状、高斯3种模型有效的描述,空间相关性存在的尺度范围变异大,中等和强空间相关性的影响范围在15—155m之间,10块样地LAI的空间相关性的影响范围平均为65.637m。普通克里格插值结果显示,LAI的空间分布呈明显的条带状和斑块状的梯度变化。LAI与林分平均胸径、林分平均高和林分平均冠长呈显著负相关,与林分平均冠幅、林分密度以及树种个数呈显著正相关。研究结果可为不同尺度云冷杉针阔混交林LAI的估计提供依据。     

广西桉树人工林叶面积指数的估测及其校正

叶面积指数(Leaf area index, LAI)是森林生态系统重要的结构参数,通过遥感技术可反演区域LAI,但其可靠性需要地面准确的实测数据进行验证。选取广西国有高峰林场不同林龄的桉树(Eucalyptus robusta)人工林为对象,以异速生长法(Allometry)为对照,综合利用植物冠层分析仪法(LAI-2200)、跟踪辐射和冠层结构分析仪法(TRAC)、半球摄影法(DHP)以及地基激光雷达法(TLS)等间接法估测样地的LAI,并考虑木质成分以及聚集效应影响,进行相应的校正处理,为地面快速、准确测量桉树人工林LAI提供参考。结果表明：桉树人工林的比叶面积为125.37±13.38 cm~2/g,通过Allometry获得的LAI变化范围在1.65—3.84,平均为2.73,不同林龄间的差异均显著(P<0.05),随着林龄的增加呈现先增加后减少的趋势。在未校正情况下,LAI-2200、TRAC、DHP、TLS估算的LAI存在显著差异(P<0.05)。与对照相比,LAI-2200在幼龄林和过熟林中估算误差最小,TRAC在成熟林中估算误差最小。相对于完全去除法,利用...     

Direct and indirect estimates of Leaf Area Index (LAI) for lodgepole and loblolly pine stands

We compared direct and indirect estimates of leaf area index (LAI) for lodgepole and loblolly pine stands. Indirect estimates of LAI using radiative methods of the LI-COR LAI-2000 Plant Canopy Analyzer (PCA) did not correlate with allometric estimates for lodgepole pine, and correlated only weakly with litter-trap estimates for loblolly pine. The PCA consistently under-estimated LAI in lodgepole pine stands with high LAI, and over-estimated LAI in the loblolly pine stands with low LAI. We developed a physical model to test the hypothesis that the PCA may under-estimate LAI in high leaf area stands because of increased foliage overlap and, therefore, increased selfshading. Radiative estimates of LAI using the PCA for the physical model were consistenly lower than allometric measures. Results from the physical model suggested that increased foliage overlap decreased the ability of the PCA to accurately estimate LAI. The relationship between allometric and radiative measures suggested an upper asymptote in LAI estimated using the PCA. The PCA may not accurately estimate LAI in stands of low or high leaf area index, and the bias or error associated with these estimates probably depends on species and canopy structure. A species specific correction factor will not necessarily correct bias in LAI estimates using the PCA.     

Altitudinal Change in LAI and Stand Leaf Biomass in Tropical Montane Forests: a Transect Study in Ecuador and a Pan-Tropical Meta-Analysis

Abstract Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm² g⁻¹ whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.     

Canopy development of a model herbaceous community exposed to elevated atmospheric CO2 and soil nutrients

To test the prediction that elevated CO₂ increases the maximum leaf area index (LAI) through a stimulation of photosynthesis, we exposed model herbaceous communities to two levels of CO₂ crossed with two levels of soil fertility. Elevated CO₂ stimulated the initial rate of canopy development and increased cumulative LAI integrated over the growth period, but it had no effect on the maximum LAI. In contrast to CO₂, increased soil nutrient availability caused a substantial increase in maximum LAI. Elevated CO₂ caused a slight increase in leaf area and nitrogen allocated to upper canopy layers and may have stimulated leaf turnover deep in the canopy. Gas exchange measurements of intact communities made near the time of maximum LAI indicated that soil nutrient availability, but not CO₂ enrichment, caused a substantial stimulation of net ecosystem carbon exchange. These data do not support our prediction of a higher maximum LAI by elevated CO₂ because the initial stimulation of LAI diminished by the end of the growth period. However, early in development, leaf area and carbon assimilation of communities may have been greatly enhanced. These results suggest that the rate of canopy development in annual communities may be accelerated with future increases in atmospheric CO₂ but that maximum LAI is set by soil fertility.     

Seasonal variability in the effect of elevated CO2 on ecosystem leaf area index in a scrub-oak ecosystem

We report effects of elevated atmospheric CO₂ concentration (C_a) on leaf area index (LAI) of a Florida scrub‐oak ecosystem, which had regenerated after fire for between three and five years in open‐top chambers (OTCs) and was yet to reach canopy closure. LAI was measured using four nondestructive methods, calibrated and tested in experiments performed in calibration plots near the OTCs. The four methods were: PAR transmission through the canopy, normalized difference vegetation index (NDVI), hemispherical photography, and allometric relationships between plant stem diameter and plant leaf area. Calibration experiments showed: (1) Leaf area index could be accurately determined from either PAR transmission through the canopy or hemispherical photography. For LAI determined from PAR transmission through the canopy, ecosystem light extinction coefficient (k) varied with season and was best described as a function of PAR transmission through the canopy. (2) A negative exponential function described the relationship between NDVI and LAI; (3) Allometric relationships overestimated LAI. Throughout the two years of this study, LAI was always higher in elevated C_a, rising from, 20% during winter, to 55% during summer. This seasonality was driven by a more rapid development of leaf area during the spring and a relatively greater loss of leaf area during the winter, in elevated C_a. For this scrub‐oak ecosystem prior to canopy closure, increased leaf area was an indirect mechanism by which ecosystem C uptake and canopy N content were increased in elevated C_a. In addition, increased LAI decreased potential reductions in canopy transpiration from decreases in stomatal conductance in elevated C_a. These findings have important implications for biogeochemical cycles of C, N and H₂O in woody ecosystems regenerating from disturbance in elevated C_a.     

利用凋落物法和林木因子模拟小兴安岭阔叶红松林叶面积指数

叶面积指数(LAI)是研究森林生态系统生理生态进程中关键的结构参数之一。目前,凋落物法是在非破坏性条件下能直接测定森林生态系统LAI的最有效的方法,然而将凋落叶按树种分类增加了该方法的实施难度。平均优势度模型、林分优势度模型和局域优势度模型基于凋落物法和林木因子(如胸高断面积basal area,BA;坐标)能精确地预测落叶阔叶林的LAI,而这些模型是否适用于针阔混交林仍未进行验证。以小兴安岭阔叶红松林(Pinus koraiensis)为研究对象,先利用凋落物法测定其LAI,依此为参考对3种模型预测LAI的有效性进行验证,并以红松、冷杉(Abies nephrolepis)、紫椴(Tilia amurensis)、五角槭(Acer mono)、枫桦(Betula costata)和裂叶榆(Ulmus laciniata)为例,探讨了基于凋落物法测定的LAI与BA的相关关系。结果表明:平均优势度模型不适于预测针阔混交林的LAI;林分优势度模型预测效果较好,精度达86%;局域优势度模型预测效果最优,精度高于90%。然而,为准确测定阔叶红松林的LAI,应最少选择测定8个主要树种的比叶面积。基于凋落物法测定的6个树种的LAI与其BA均显著相关(P0.01),最小R2为0.67。研究结果可为快速、准确地测定针阔混交林的LAI提供依据,为非破坏性条件下建立树种的LAI与其BA的相关关系提供参考。     

基于不同空间尺度遥感影像估算森林叶面积指数的差异

地面测量点对遥感像元的代表性如何,怎样获取像元的相对真值,多大的空间分辨率可以真实地反映森林区域的叶面积指数(LAI),这些都是定量遥感中的重要问题.本研究计算LAI-2200和TRAC两种冠层分析仪测量的空间范围,并结合GF-2(4.1 m)、Sentinel-2(10 m)、Landsat-8(30 m)3种不同空间分辨率遥感影像,找到各尺度下像元的相对真值,在保持真值观测面积和遥感获取面积一致的条件下,基于一元指数和多元回归模型,对比分析不同空间分辨率影像对估算森林LAI的影响,并对3种影像模型进行30和100 m尺度下的检验以及各自数据集的空间代表性评价,比较得出最适合表达研究区域森林LAI的尺度.结果表明:对于森林来说,高分辨率并不一定能充分反映森林LAI.基于3种分辨率影像的统计模型都能很好地估测森林LAI,其中,基于Sentinel-2的反演精度最高,基于GF-2的反演精度最低.30和100 m尺度下的检验结果表明,基于GF-2反演模型高估了森林LAI,基于Landsat-8的反演模型低估了森林LAI,基于Sentinel-2分辨率的统计模型可以很好地估测研究区域森林LAI.     

Implications of shoot structure on the rate of photosynthesis at different levels in a coniferous canopy using a model incorporating grouping and penumbra

1. The connection between high leaf area index (LAI) and photosynthetic production with two attributes of coniferous canopy structure: small leaf size and grouping of needles on shoots, was analysed using a simulation model.
2. The small size of conifer needles gives rise to penumbras, which even out the distribution of direct sunlight on the leaf area and thereby act to increase the rate of canopy photosynthesis per unit of LAI.
3. Grouping, by producing a non-uniform distribution of leaf area, causes a decrease in total canopy light interception at any given LAI, but improves the photosynthetic light capture by shoots in the lower canopy.
4. Application of the model on a case study showed that: (a) grouping had a negative effect on the rate of photosynthesis in the upper canopy, but deeper down in the canopy the situation was reversed; (b) in the lower canopy, photosynthetic rates were up to 10 times higher as a result from the combined effect of grouping and penumbra; (c) grouping did not improve the rate of canopy photosynthesis per unit of LAI, however, it can have a positive effect on the total photosynthetic production by allowing a higher productive LAI to be maintained; (d) penumbra, on the other hand, increased the rate of canopy photosynthesis by as much as 40% for moderate values of the LAI.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

基于多角度植被指数的马尾松林LAI反演方法

CHRIS/PROBA是目前具有最高空间分辨率（17 m×17 m）的星载多角度高光谱数据,该款数据在反演植被垂直结构参数,如树高、叶面积指数（leaf area index,LAI）等方面具有重要的应用前景。基于四尺度几何光学模型得到马尾松（Pinus massoniana Lamb.）冠层的归一化差分植被指数（normalized difference vegetation index,NDVI）各向异性分布规律,利用CHRIS红光特征波段和近红外特征波段构建一种新型多角度植被指数（normalized hotspot-dark-spot difference vegetation index,NHDVI）,并将其应用于CHRIS数据对马尾松林的LAI遥感估算上。结果显示：（1）相比归一化差分植被指数（NDVI）与土壤调节植被指数（soil adjusted vegetation index,SAVI）而言,NHDVI能很好地融合光谱信息与角度信息,与地面实测LAI的决定系数达到0.7278;（2）利用NHDVI-LAI统计回归模型方法来反演LAI值,将得到的LAI值与地面实测值进行相关性分析,结果拟合优度达到0.8272,均方根误差RMSE为0.1232。与传统植被指数相比,包含角度信息的多角度植被指数对LAI的反演在精度上有较大提升,同时比基于辐射传输模型的反演方法更简易、实用。     

Factors contributing to accuracy in the estimation of the woody canopy leaf area density profile using 3D portable lidar imaging

Factors that contribute to the accuracy of estimating woody canopy's leaf area density (LAD) using 3D portable lidar imaging were investigated. The 3D point cloud data for a Japanese zelkova canopy [Zelkova serrata (Thunberg) Makino] were collected using a portable scanning lidar from several points established on the ground and at 10 m above the ground. The LAD profiles were computed using voxel-based canopy profiling (VCP). The best LAD results [a root-mean-square error (RMSE) of 0.21 m(2) m(-3)] for the measurement plot (corresponding to an absolute LAI error of 9.5%) were obtained by compositing the ground-level and 10 m measurements. The factors that most strongly affected estimation accuracy included the presence of non-photosynthetic tissues, distribution of leaf inclination angles, number (N) of incident laser beams in each region within the canopy, and G(theta(m)) (the mean projection of a unit leaf area on a plane perpendicular to the direction of the laser beam at the measurement zenith angle of theta(m)). The influences of non-photosynthetic tissues and leaf inclination angle on the estimates amounted to 4.2-32.7% and 7.2-94.2%, respectively. The RMSE of the LAD estimations was expressed using a function of N and G(theta(m)).