上层冠层因子对马尾松肉桂复层林下层林分生物量的影响

采用标准地定位监测方法,选取不同栽培模式马尾松(Pinus massoniana)肉桂(Cinnamomum cassia)人工复层林上层林冠的叶面积指数、平均叶倾角、直接辐射透过系数、散射辐射透过系数、消光系数等5个冠层因子作为影响下层林分生长的光环境指标,利用主成分分析法研究影响肉桂林分生物量的主要光环境影响因子。结果表明,不同上层密度光环境下的肉桂各器官生物量变化明显;在主成分分析中,马尾松冠层叶面积指数、平均叶倾角和直接辐射透过系数三者的方差贡献率累计达到99.37%,是影响下层林生长的主导光环境因子;肉桂生物量的主成分回归模型具有良好的拟合度,从回归模型中可得出影响肉桂生物量的主要因子是上层林冠层的直接辐射透过系数。     

森林冠层叶片氮浓度高光谱遥感反演研究进展

氮素是植物生命活动所需的重要营养元素，在森林植被的光合作用和生态系统固碳方面起着关键作用。因此，理解森林叶片氮浓度在叶片和冠层（遥感像元）尺度上的高光谱特征，是开展森林冠层叶片氮浓度（CNC）遥感反演、优化森林碳循环模拟、应对气候变化的重要基础工作。当前，森林CNC的光谱特征提取受到冠层结构因素的影响，其高光谱遥感反演的理论亦不明确。文中通过梳理国内外大量植被叶氮高光谱反演的代表性研究成果，以时间为轴线从叶片和冠层2个尺度上进行文献综述，详细阐述当前国内外森林叶氮浓度高光谱遥感反演的主要方法、研究热点和面临的问题，并对近年来学界关于森林冠层结构在冠层叶片氮浓度遥感反演中的影响进行综述，并展望森林冠层叶氮浓度高光谱遥感反演的发展方向。     

冠层结构对林下经济植物的影响

应用美国Co-operatively公司生产的DELTA-T冠层分析仪及软件,对辽东地区5种林下经济植物生长区域林下光环境特性和冠层结构指标光合光量子通量密度(PPFD)、总空隙度(gap fraction)、叶面积指数(LAI)、平均叶倾角(MLA)等的研究结果表明,大叶芹和五味子更喜欢林下直射光,而移山参、穿龙薯蓣和细辛更适应林下散射光。     

毛竹枝叶生物量的冠层分布对钩梢和施肥的响应

以浙江省不钩梢和钩梢2种不同营林模式下的毛竹Phyllostachys heterocycla cv.pubescens为研究对象,并设置施肥处理,对林地毛竹枝、叶生物量在冠层的分布进行了研究。结果表明:不钩梢毛竹枝、叶和枝叶总生物量在冠层上的分布符合Peel-Reed模型,而钩梢毛竹符合二次曲线分布;施肥增加毛竹枝、叶生物量,在钩梢与不钩梢之间存在差异,前者在各枝盘上,枝生物量的增加均大于叶,而后者在主要枝盘(≥8枝盘)上枝生物量的增加则小于叶;毛竹叶枝比在钩梢或不钩梢毛竹上总体表现为随着枝盘数的增加而递增,但钩梢毛竹在所有枝盘上的叶枝比均小于1;施肥表现为相对地增大了冠层中下部(如不钩梢23~30枝盘和钩梢1~9枝盘)的叶枝比;钩梢(剩余14枝盘)毛竹叶片生物量仅为不钩梢毛竹的58.99%,且树冠形状呈倒三角形,不利于冠层叶片充分利用光资源;施肥在一定程度可改善树冠形状,但生产上应适度降低毛竹钩梢强度,保留枝盘数为18~20枝盘,有效保持或提升毛竹叶片生物量。     

辽东栎冠层叶建成消耗与比叶面积的空间异质性

植物功能性状是目前生态学领域关注的热点问题之一,而比叶面积(Specific Leaf Area,SLA)和叶建成消耗(Leaf Construction Cost,LCC)则是功能性状中研究植物资源收获与支出的首选指标。树冠冠层结构与功能在方向上的不同可导致冠层功能性状空间分布的差异。以东灵山辽东栎为研究对象,分析了不同冠层不同方向部位叶片的功能性状以及影响因素。结果表明:在不同空间上,冠层上部的CC均高于冠层底部的,而SLA则相反;除上部与东向外,各方向叶片的比叶面积均有显著差异;热值则是影响二者的主要因素。冠层CC和SLA的空间异质性研究,对于在冠层水平上揭示植物资源利用具有重要意义。     

兴安落叶松人工林冠层气体交换的时空特性

在生长季定期测定兴安落叶松林主要组成树种兴安落叶松和白桦树冠上部、中部和下部叶片的光合作用和呼吸作用.结果表明:兴安落叶松当年生枝针叶在8、9月份最大净光合速率在冠层间差异显著.8月份最高,从上至下分别为13.09、8.63和9.07 μmolCO2·m-2s-1.落叶松往年生枝针叶的最大净光合速率在生长季差异不显著,其平均最大净光合速率为8.22 μmolCO2·m-2s-1.在整个生长季白桦冠层间的最大净光合速率均差异显著,从上至下,依次为17.56、13.01和9.58 μmolCO2·m-2s-1;白桦叶片、落叶松当年生枝针叶和落叶松往年生枝针叶的暗呼吸速率差异随着时间的变化而发生改变,但冠层之间无显著差异.在整个生长季中,白桦叶片、落叶松当年生枝针叶和落叶松往年生枝针叶的暗呼吸速率分别为0.47～1.40,0.83～1.46和0.68～1.63 μmolCO2·m-2s-1.2树种的暗呼吸速率与叶片温度显著正相关,与光照不相关.除白桦冠层上层叶片外,光合作用与生长季叶片温度的变化无明显相关性;白桦和落叶松生长季的不同时期冠层气体交换是动态变化的,但变化式样具有种的特异性.     

苏南丘陵区主要林型叶面积指数季节动态

应用LAI-2000冠层分析仪,研究了苏南丘陵区主要林型的叶面积指数(LAI)季节动态。结果表明:4种林型的LAI随季节变化具有明显的规律性,充分反映了4种林型叶片的生长状态,LAI值由大到小依次为毛竹(3.45)>杉木(2.87)>麻栎(2.62)>马尾松(2.00);冠层空隙度总体变化趋势与LAI的总体变化趋势恰好相反,在8月份出现最小值,分别为杉木6.7%、毛竹2.8%、麻栎5.6%、马尾松15.1%;叶倾角的季节变化不大,均在一定范围内波动。对4种林型的冠层均匀性进行比较分析,结果发现,杉木林冠层均匀性最好,麻栎林冠层均匀性最差。对4种林型的叶面积指数与冠层空隙度进行回归分析,发现两者之间呈指数回归关系(R2>0.95),通式为:y=ae-bx。     

黄土高原区不同密度刺槐林冠层结构特征及月动态变化

为合理经营黄土高原区刺槐人工林,利用LAI-2200植物冠层分析仪,研究了不同密度刺槐人工林冠层结构特征及月变化。结果表明:密度对LAI(叶面积指数)有影响,随着密度的增加,林分叶面积指数趋于一致;在生长月份上,LAI为先增后降趋势,最大值出现在6月底。密度与冠层开度呈显著负相关,随着密度的增加,刺槐林DIFN明显降低,925～1 125株/hm2与其他3个密度林分有显著性差异;DIFN(无截取散射)随月份的变化均呈"V"字型变化,在6月底至8月初DIFN最低。密度与MTA(平均叶倾角)呈显著正相关,密度在925～1 125株/hm2范围的刺槐林分,叶片几乎处于水平状态,其他3种密度林分MTA均在40°～50°之间有最大分布频率,而月份对MTA影响不显著。对叶面积指数与冠层开度进行回归分析,发现两者之间呈指数回归关系(R2=0.998)。     

上海人工绿地群落UVB屏蔽效率与冠层特征关系初步研究

采用欧氏距离聚类分析方法,对上海几种典型植物群落的UVB屏蔽效率和群落叶面积指数、枝下高等冠层特征因子日的相关关系进行分析。结果表明：1）所测定人工群落UVB平均屏蔽效率为91．1％,群落内2m高处UVB辐射通量仅为全光照旷地的14％。毛竹群落和女贞群落屏蔽效率最高,分别为98．5％,97．8％。2）不同植物群落UVB屏蔽效率在冠层特征梯度上表现出显著的分异,植物群落内UVB辐射通量与群落叶面积指数成显著负相关关系,与群落高度和枝下高成显著正相关关系。3）在植被覆盖较少且紫外地面反射辐射较为强烈的游憩场所,可考虑种植叶倾角较小且叶面积较大的植物,如广玉兰（Magnolia grandiflora）、女贞（Ligustmm lucidum）、桂花（Osmanthus fragrans）等。     

雷竹林冠层特性与叶片的空间分布

研究了雷竹林的冠层特性和叶片的空间分布规律。结果表明 ,随着雷竹竹冠部位上升 ,分枝角度逐渐变小 ,而立竹密度变大 ,分枝角度略偏大 ;竹株不同留枝数量、胸径和林分不同立竹密度对叶量分布有较大影响 ,随着留枝盘数的增加 ,单株竹株的全叶重显著增加 ,在留枝盘数为 1 5～ 1 6档时 ,每株叶量最大 ;随胸径增大 ,全株叶重增大 ,在胸径为 3 7cm左右时叶量增幅最小 ,而后趋于平缓 ;立竹密度的增加 ,导致林冠叶量重心逐渐上移 ,立竹密度为 2 5 6～ 2 92× 1 0 4株·hm-2 ,林冠上层叶重比例达到 6 3 1 3%,林冠下层自然落叶现象明显。     

日本落叶松冠层光合特性的空间变化

【目的】研究日本落叶松冠层间光合特性的差异,揭示冠层光合特性的空间变化规律,为精确估算冠层生产力及构建冠层生产力模型提供依据。【方法】以8年生日本落叶松为研究材料,根据树冠长度,将树冠等分为8个层级,从上到下分别为第1~8层级,选择每一层级的南向一级侧枝作为光合测定枝条,测定每一枝条上针叶的光合日变化。【结果】多数冠层间光合参数日变化规律相似,净光合速率、蒸腾速率、气孔限制值、光合有效辐射、气温和水汽压亏缺均表现为单峰曲线,气孔导度从上午开始逐渐降低。随着冠层的升高,光合参数增加、比叶面积减小,而气温和水汽压亏缺表现为先增后降的趋势,不同冠层间均差异显著( P <0.05)。净光合速率、光合有效辐射、气孔导度和蒸腾速率与相对冠高存在指数函数关系( R^2 ≥0.94),气温和水汽压亏缺与相对冠高存在一元二次函数关系(R^2 ≥0.79)。通径分析表明,影响净光合速率的主要因素是光合有效辐射,而气温和空气相对湿度影响较小。【结论】光合有效辐射和叶肉细胞的光合活性分别是影响日本落叶松净光合速率空间变化的主要环境因素和生理因素。净光合速率随冠层的升高而显著增加,而冠层底部净光合速率为负值,始终处于碳消耗状态。     

Analysis of the sensitivity of absorbed light and incident light profile to various canopy architecture and stand conditions

We analyzed the effect of simplifying assumptions in canopy representation of radiation transfer models, comparing modeled diffuse non-interceptance and photosynthetic photon flux density with measurements at different layers of complex pine-broadleaved canopy with large seasonal variation of leaf area index. The most detailed model included clumping of trees (i.e.,?stand density) and a vertical specification of leaf angle distribution and shoot clumping. A less detailed model replaced the vertically specified variables with their means. The most parsimonious model accounted for neither shoot clumping nor stand density. The vertical specification of shoot clumping and leaf angle distribution only slightly improved vertical and seasonal openness and light estimates over using mean values. Further simplification had little effect on total absorbed light but was more risky for estimates of the vertical distributions of openness and light absorbed by the canopy, which will affect photosynthesis estimates due to the non-linearity of photosynthetic light response. Including woody surfaces in winter, when leaf area was low, was essential for reproducing the measurements correctly. A sensitivity analysis showed that ignoring (i)?shoot clumping could result in a substantial overestimation of total absorbed light with errors increasing with decreasing leaf area and (ii) stand density in sparse stands could lead to substantial overestimation of total absorbed light, and the effect is largely independent of leaf area. Also, (iii) the effect of changing leaf angle distribution increased with decreasing leaf area, and was larger and more persistent along the leaf area range with increasing shoot clumping. Overall, accounting for the effect of tree clumping on absorbed light is most important in stands composed of species where leaves are not very clumped (e.g., broadleaved). However, even in forests with highly clumped shoots (i.e., coniferous), an accurate estimation of absorbed light distribution in stands requires incorporation of stand density in the model.     

Importance of needle age and shoot structure on canopy net photosynthesis of balsam fir (Abies balsamea): a spatially inexplicit modeling analysis

We have developed a spatially inexplicit model of canopy photosynthesis for balsam fir (Abies balsamea (L.) Mill.) that accounts for key processes of light-shoot interaction including irradiance interception by the shoot, spatial aggregation of shoots into branches and crowns, the differential propagation of diffuse and direct light within the canopy, and an ideal representation of penumbra. Also accounted for in the model are the effects of the average radiative climate and shoot age on needle retention, light interception, and photosynthetic capacity. We used reduced versions of this model to quantify the effects of simplifying canopy representation on modeled canopy net photosynthesis. Simplifications explored were the omission of direct beam transformation into penumbral light and the use of different constant shoot properties throughout the canopy. The model was parameterized for a relatively dense balsam fir stand (leaf area index of 5.8) north of Québec City, Canada, and run using hourly meteorological data obtained at the site. The overall performance of the complete model was satisfactory, with maximum values of canopy net photosynthesis of 23 micromol (m(2) ground)(-1) s(-1) (83 mmol m(-2) h(-1)), and a near-saturation of the canopy at a photosynthetically active radiation photon flux density of about 750 micromol m(-2) s(-1) (2.7 mol m(-2) h(-1)). The omission of penumbral effects through the use of unattenuated direct (beam) radiation at all layers of the canopy, as used for broad-leaved species, reduced canopy net photosynthesis by 3.7%. Analysis of the results show that the small impact of penumbra on canopy net photosynthesis stems from the high proportion of diffuse radiation (73%) estimated from our meteorological data set; single-hour results under clear sky conditions approach theoretical bias values of about 30%. Use of mean shoot photosynthetic, light capture and light transmission properties throughout the canopy biased canopy net photosynthesis by less than 3%. However, simulations carried out based on properties of 1-year-old shoots throughout the canopy overestimated canopy net photosynthesis by 9%. Use of the shoot as our smallest functional unit was a potential source of bias because the differential absorption of direct and diffuse radiation within the shoot could not be factored into the model. Other sources of potential bias are discussed.     

Three-dimensional lamina architecture alters light-harvesting efficiency in Fagus: a leaf-scale analysis

Modification of foliage exposition and morphology by seasonal average integrated quantum flux density (Qint) was investigated in the canopies of the shade-tolerant late-successional deciduous tree species Fagus orientalis Lipsky and Fagus sylvatica L. Because the leaves were not entirely flat anywhere in the canopy, the leaf lamina was considered to be three-dimensional and characterized by the cross-sectional angle between the leaf halves (theta). Both branch and lamina inclination angles with respect to the horizontal scaled positively with irradiance in the canopy, allowing light to penetrate to deeper canopy horizons. Lamina cross-sectional angle varied from 170 degrees in the most shaded leaves to 90-100 degrees in leaves in the top of the canopy. Thus, the degree of leaf rolling increased with increasing Qint, further reducing the light-interception efficiency of the upper-canopy leaves. Simulations of the dependence of foliage light-interception efficiency on theta demonstrated that decreases in theta primarily reduce the interception efficiency of direct irradiance, but that diffuse irradiance was equally efficiently intercepted over the entire range of theta values in our study. Despite strong alteration in foliage light-harvesting capacity within the canopy and greater transmittance of the upper crown compared with the lower canopy, mean incident irradiances varied more than 20-fold within the canopy, indicating inherent limitations in light partitioning within the canopy. This extensive canopy light gradient was paralleled by plastic changes in foliar structure and chemistry. Leaf dry mass per unit area varied 3-4-fold between the canopy top and bottom, providing an important means of scaling foliage nitrogen contents and photosynthetic capacity per unit area with Qint. Although leaf structure versus light relationships were qualitatively similar in all cases, there were important tree-to-tree and species-to-species variations, as well as evidence of differences in investments in structural compounds within the leaf lamina, possibly in response to contrasting leaf water availability in different trees.     

An estimate of the photosynthetic production of individual trees in a Chamaecyparis obtusa plantation

A Weibull function was used to model the vertical distribution of leaf area of individual trees in a 25-year-old Chamaecyparis obtusa (Siebold & Zucc.) Endl. plantation. The parameter representing the shape of the leaf distribution was independent of tree size. A scale parameter tended to decrease with tree size suggesting a critical minimum height for retention of foliage by trees. On the basis of leaf distribution, the photosynthetic production of individual trees was estimated from the canopy photosynthetic production, which was determined from a model of canopy photosynthesis. The data indicated that the photosynthesis of a tree was proportional to the corresponding tree weight to the power of 1.84. Furthermore, the photosynthetic production varied as the 3/2nd power of total leaf area of the tree. Thus, it was concluded that the photosynthetic production per unit of leaf area, that is, the mean photosynthetic activity of a tree, is proportional to the stem girth at clear length, or the square root of the leaf area of the tree.     

Canopy development in young sugi (Cryptomeria japonica) stands in relation to changes with age in crown morphology and structure

In young sugi (Cryptomeria japonica D. Don) stands, crown shape (crown length/crown diameter) ratio, average branch inclination, and spatial density of foliage in the crown increased with stand age. Within crowns, foliage distribution increased from the apex downward and, until crown closure, reached a maximum near the crown base. After crown closure, the maximum occurred near the middle of the crown. In each stand, foliage distribution in the canopy showed almost the same vertical change over time as it did in individual crowns. The vertical distribution of foliage in the canopy moved upward with stand age, accompanied by an increase in canopy depth and leaf mass. The shape of the vertical distribution was almost symmetrical between the upper and lower halves in the closed stands, although slightly skewed downward. The logarithm of average spatial density decreased linearly as cumulative leaf mass increased with distance from the top of the canopy. The total cross-sectional area of the crowns exceeded the stand area from the middle of the canopy downward in the closed stands because of crown overlap. However, partly because of changes in crown morphology and structure, the increase in leaf mass with stand age did not always cause more severe crown competition.     

基于简单竞争指数的杉木人工林树冠形状模拟

以湖南省株洲市攸县黄丰桥国有林场杉木中龄林为研究对象,用逐步回归的方法建立以单木大小、单木竞争指标为自变量,冠幅、冠长为因变量的函数估算模型,实现林木树冠形状的模拟。研究结果显示:在杉木树冠形状模拟时,与竞争有关的冠幅模型、冠长模型的判定系数R2分别为0.771、0.872,显著高于与竞争无关的冠幅模型、冠长模型的调整的判定系数0.729、0.811,这说明竞争对树冠生长影响的作用是显著的。     

Summary models for light interception and light-use efficiency of non-homogeneous canopies

The application of detailed models of canopy photosynthesis rely on the estimation of attenuation of light in the canopy. This attenuation is readily estimated with the Lambert-Beer law when the canopy is homogeneous. In reality, forest canopies are far from homogeneous, and this has led to the use of detailed light extinction models that account for grouping of foliage between and within trees. Because such models require detailed parameterization and fine resolution inputs, they are impractical in larger-scale applications. Thus, there is interest in simplified models that can be readily parameterized. We developed two equations that can be used to estimate mean annual light interception by single unshaded trees and by stands of Poisson distributed trees. Interception by single trees is a function of crown surface area, the ratio of leaf area to crown surface area, the extinction coefficient in a homogeneous canopy--which can be determined separately--and one empirical parameter that depends on the mean solar angle. The summary model was tested against a detailed model of interception, and showed good agreement, although with slight bias. The results showed that crown surface area is a good summary variable for crown size and shape, because errors are independent of crown shape (ellipsoids, cones and height:width ratios). We also tested whether canopy photosynthesis is proportional to light interception across canopies differing in structure and leaf area index, and found that light-use efficiency is influenced by canopy structure. The model is useful in larger-scale applications because it can be parameterized with available data without the need for additional empirical parameters. It can also be used to study the effect of stand structure on mean annual light interception and productivity.     

Modeling radiation transfer within the canopy of a Chinese fir plantation

Calculation of radiation transfer within the canopy of a plantation on sloped sites is described. The canopy was assumed to consist of evenly cone-shaped crowns. Within-tree shading by object tree and between-tree shading by tree crowns around the object tree were differentiated. The two-dimensional needle area density (NAD) was introduced in the calculation of radiation transfer and interception. The model was validated with measurements of both daily photosynthetic photo flux density (PPFD) and daily photosynthetic radiation at tens of points within the crown of a 17-year-old Chinese fir (Cunninghamia lanceolata) plantation growing on a sloped site. Simulation showed that modeled data closely matched measurements. Simulation errors likely result from the uneven size of tree crowns, irregular crown shape, grouping of needles to shoots and twigs, etc. Scattering radiation amounted to below 10%, in most cases, of daily radiation, and its uncertainty was relatively small.     

Leaf area distribution and radiative transfer in open-canopy forests: implications for mass and energy exchange

Leaf area and its spatial distribution are key canopy parameters needed to model the radiation regime within a forest and to compute the mass and energy exchange between a forest and the atmosphere. A much larger proportion of available net radiation is received at the forest floor in open-canopy forests than in closed-canopy forests. The proportion of ecosystem water vapor exchange (lambda E) and sensible heat exchange from the forest floor is therefore expected to be larger in open-canopy forests than in closed-canopy forests. We used a combination of optical and canopy geometry measurements, and robust one- and three-dimensional models to evaluate the influence of canopy architecture and radiative transfer on estimates of carbon, water and energy exchange of a ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forest. Three-dimensional model simulations showed that the average probability of diffuse and direct radiation transmittance to the forest floor was greater than if a random distribution of foliage had been assumed. Direct and diffuse radiation transmittance to the forest floor was 28 and 39%, respectively, in the three-dimensional model simulations versus 23 and 31%, respectively, in the one-dimensional model simulations. The assumption of randomly distributed foliage versus inclusion of clumping factors in a one-dimensional, multi-layer biosphere-atmosphere gas exchange model (CANVEG) had the greatest effect on simulated annual net ecosystem exchange (NEE) and soil evaporation. Assuming random distribution, NEE was 41% lower, net photosynthesis 3% lower, total lambda E 10% lower, and soil evaporation 40% lower. The same comparisons at LAI 5 showed a similar effect on annual NEE estimates (37%) and lambda E (12%), but a much larger effect on net photosynthesis (20%), suggesting that, at low LAI, canopies are mostly sunlit, so that redistribution of light has little effect on net photosynthesis, whereas the effect on net photosynthesis is much greater at high LAIs.