基于Campbell椭球分布函数的大兴安岭地区主要树种叶倾角分布模拟

叶倾角分布(LAD)直接决定着植被冠层对辐射的截获量,同时对入射太阳辐射的大小与走向也起着决定性作用,是定量遥感中的关键参数.本研究基于Campbell椭球分布函数和迭代方法拟合大兴安岭林区主要树种的LAD,定量分析冠层叶片分层与不分层时模型的拟合情况及不同龄组对LAD的影响.结果表明： 大兴安岭地区6种主要树种的LAD均属于横椭球分布,针叶树的平均叶倾角小于阔叶树;无论对叶片分层处理与否,模型拟合叶倾角的结果与实测结果基本一致;白桦和落叶松的拟合结果与实测结果线性回归的相关系数分别是0.8268、0.8192,均方根误差分别是3.7%、4.3%,说明Campbell模型应用于森林冠层是可靠的;考虑龄组的影响时,虽然分层处理时叶倾角的分布趋势与龄组无关,但幼龄落叶松的平均叶倾角小于成熟落叶松,表明龄组对叶倾角分布取值有正向影响,而对消光系数取值有负向
影响.     

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。     

Effect of light interception by non-photosynthetic organs on canopy photosynthetic production

A canopy photosynthesis model was derived on the assumption that the light diminution within a canopy is caused by both leaves and non-photosynthetic organs. The light diminution by leaves and that by non-photosynthetic organs were taken into account separately in the Lambert-Beer equation of light extinction. The light flux density on the leaf surface at each depth was evaluated from the leaf's share of light. The light flux density on the leaf surface thus obtained was incorporated into the Monsi-Saeki model of canopy photosynthesis. The proposed model was applied for estimating gross canopy photosynthesis in a 19-year-oldLarix leptolepis plantation where 38% of the light diminution was due to non-photosynthetic organs. The daily canopy photosynthesis on one summer day calculated using the present model was about 22% less than that calculated by the conventional Monsi-Saeki model, in which light interception by non-photosynthetic organs is neglected. The degree of such reduction in canopy photosynthesis through shading by non-photosynthetic organs was assessed in relation to parameters affecting light extinction, leaf photosynthetic characteristics, and light regime above the canopy.     

基于PROSAIL辐射传输模型的毛竹林叶面积指数遥感反演

采用PROSAIL辐射传输模型建立毛竹林叶面积指数(LAI) 冠层反射率查找表,并结合Landsat TM卫星遥感数据,实现了毛竹林LAI的定量反演.结果表明： PROSAIL模型各输入参数的敏感性由高到低依次为LAI>叶绿素含量(C_ab)>叶片结构参数(N)>平均叶倾角(ALA)>等效水厚度(C_w)>干物质含量(C_m),并以LAI、C_ab两个主要敏感因子用于构建毛竹林LAI 冠层反射率查找表;基于PROSAIL模型的毛竹林LAI遥感反演结果与实测LAI具有很好的一致性,二者相关系数为0.90,均方根误差和相关的均方根误差也较小,分别为0.58和13.0%,但也存在反演LAI平均值高于实际值的问题.     

Harnessing Genetic Variation in Leaf Angle to Increase Productivity of Sorghum bicolor

The efficiency with which a plant intercepts solar radiation is determined primarily by its architecture. Understanding the genetic regulation of plant architecture and how changes in architecture affect performance can be used to improve plant productivity. Leaf inclination angle, the angle at which a leaf emerges with respect to the stem, is a feature of plant architecture that influences how a plant canopy intercepts solar radiation. Here we identify extensive genetic variation for leaf inclination angle in the crop plant Sorghum bicolor, a C4 grass species used for the production of grain, forage, and bioenergy. Multiple genetic loci that regulate leaf inclination angle were identified in recombinant inbred line populations of grain and bioenergy sorghum. Alleles of sorghum dwarf-3, a gene encoding a P-glycoprotein involved in polar auxin transport, are shown to change leaf inclination angle by up to 34° (0.59 rad). The impact of heritable variation in leaf inclination angle on light interception in sorghum canopies was assessed using functional-structural plant models and field experiments. Smaller leaf inclination angles caused solar radiation to penetrate deeper into the canopy, and the resulting redistribution of light is predicted to increase the biomass yield potential of bioenergy sorghum by at least 3%. These results show that sorghum leaf angle is a heritable trait regulated by multiple loci and that genetic variation in leaf angle can be used to modify plant architecture to improve sorghum crop performance.     

双季稻光合生产模型的建立与应用

综合已有作物模型(包括冠层结构、冠层光分布和冠层光合作用与干物质生产模型)的优点,构建了双季稻光合生产模型.利用独立的田间试验资料,对冠层内的光分布和干物质积累量进行了初步检验;利用模型定量分析了直接辐射在上挺下挺、上挺下披和上披下披3种典型株型水稻冠层内水平面上和叶面上的分布、冠层日光合量及其随叶面积指数的变化特征.结果表明: 模拟值与观测值之间具有较好的一致性,预测双季稻冠层内光分布的根均方差、相对根均方差和相关系数分别为12.01 J·m^-2·s^-1、8.2%和0.9929;预测双季稻干物质积累量的根均方差、相对根均方差和相关系数分别为0.83 t·hm^-2、14.6%和0.9772,表明模型预测性较好;上挺下披株型水稻的冠层日光合量最高,取决于较大的叶面受光量、叶片光合效能和叶面积指数.     

Above-ground biomass estimation in closed canopy Neotropical forests using lidar remote sensing: factors affecting the generality of relationships

Aim Previous studies have developed strong, site‐specific relationships between canopy metrics from lidar (light detecting and ranging) remote sensing data and forest structural characteristics such as above‐ground biomass (AGBM), but the generality of these relationships is unknown. In this study, we examine the generality of relationships between lidar metrics and forest structural characteristics, including AGBM, from two study areas in Central America with different precipitation patterns. Location A series of tropical moist forest sites in Panama and a tropical wet forest in Costa Rica. Methods Canopy metrics (e.g. canopy height) were calculated from airborne lidar data. Basal area, mean stem diameter and AGBM were calculated from measurements taken as a part of ongoing forest dynamics studies in both areas. We examined the generality of relationship between lidar metrics and forest structure, and possible environmental effects (e.g. leaf phenology). Results We found that lidar metrics were strongly correlated (R²: 0.65–0.92) with mean stem diameter, basal area and AGBM in both regions. We also show that the relationships differed between these regions. Deciduousness of canopy trees in the tropical moist forest area accounted for the differences in predictive equations for stem diameter and basal area. The relationships between lidar metrics and AGBM, however, remained significantly different between the two study areas even after adjusting for leaf drop. We attribute this to significant differences in the underlying allometric relationships between stem diameter and AGBM in tropical wet and moist forests. Conclusions Important forest structural characteristics can be estimated reliably across a variety of conditions sampled in these closed‐canopy tropical forests. Environmental factors such as drought deciduousness have an important influence on these relationships. Future efforts should continue to examine climatic factors that may influence the generality of the relationships between lidar metrics and forest structural characteristics and assess more rigorously the generality of field‐derived allometric relationships.     

Complex adjustments of photosynthetic potentials and internal diffusion conductance to current and previous light availabilities and leaf age in Mediterranean evergreen species Quercus ilex

Mature non-senescent leaves of evergreen species become gradually shaded as new foliage develops and canopy expands, but the interactive effects of integrated light during leaf formation (Q(int)G), current light (Q(int)C) and leaf age on foliage photosynthetic competence are poorly understood. In Quercus ilex L., we measured the responses of leaf structural and physiological variables to Q(int)C and Q(int)G for four leaf age classes. Leaf aging resulted in increases in leaf dry mass per unit area (M(A)), and leaf dry to fresh mass ratio (D(F)) and decreases in N content per dry mass (N(M)). N content per area (N(A)) was independent of age, indicating that decreases in N(M) reflected dilution of leaf N because of accumulation of dry mass (NA = N(M) M(A)). M(A), D(F) and N(A) scaled positively with irradiance, whereas these age-specific correlations were stronger with leaf growth light than with current leaf light. Area-based maximum ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylase activity (V(cmax)A), capacity for photosynthetic electron transport (J(max)A) and the rate of non-photorespiratory respiration in light (R(d)A) were also positively associated with irradiance. Differently from leaf structural characteristics, for all data pooled, these relationships were stronger with current light with little differences among leaves of different age. Acclimation to current leaf light environment was achieved by light-dependent partitioning of N in rate-limiting proteins. Mass-based physiological activities decreased with increasing leaf age, reflecting dilution of leaf N and a larger fraction of non-photosynthetic N in older leaves. This resulted in age-dependent modification of leaf photosynthetic potentials versus N relationships. Internal diffusion conductance (g(m)) per unit area (g(m)A) increased curvilinearly with increasing irradiance for two youngest leaf age classes and was independent of light for older leaves. In contrast, g(m) per dry mass (g(m)M) was negatively associated with light in current-year leaves. Greater photosynthetic potentials and moderate changes in diffusion conductance resulted in greater internal diffusion limitations of photosynthesis in higher light. Both area- and mass-based g(m) decreased with increasing leaf age. The decrease in diffusion conductance was larger than changes in photosynthetic potentials, leading to larger CO2 drawdown from leaf internal air space to chloroplasts (delta(c)) in older leaves. The increases in diffusion limitations in older leaves and at higher light scaled with age- and light-dependent increases in MA and D(F). Overall, our study demonstrates a large potential of foliage photosynthetic acclimation to changes in leaf light environment, but also highlights enhanced structural diffusion limitations in older leaves that result from leaf structural acclimation to previous rather than to current light environment and accumulation of structural compounds with leaf age.     

冠层部位和叶龄对红松光合蒸腾特性的影响

利用Li-6400便携式CO2/H2O红外气体分析仪测定了红松不同冠层部位和叶龄针叶的光合蒸腾特性及其环境影响因子.结果表明:冠层部位和叶龄显著地影响最大净光合速率(Pmax)、光饱和点(LSP)、光补偿点(LCP)、表观最大量子效率(α)、蒸腾速率(Tr)和比叶面积(SLA),但对水分利用效率(WUE)影响不显著.随着冠层部位的下降和叶龄的增加,红松针叶的Pmax逐渐下降,其平均值变动在6·55～9·05μmol·m-2·s-1之间.不同冠层部位和叶龄针叶的LSP和LCP的差异很大,以树冠中部针叶对弱光和强光的利用能力最大.Tr随着冠层部位的下降而降低;不同叶龄针叶的Tr在1·37～1·59mmol·m-2·s-1之间变化.不同部位和叶龄红松针叶的Tr和光合有效辐射存在极显著正相关关系(R2=0·967).红松的WUE与净光合速率紧密相关(R2=0·860).随冠层部位的上升和叶龄的增大,红松针叶的SLA递减,分别在6·61～8·41m2·kg-1和6·65～8·38m2·kg-1之间波动.     

Linking light attenuation,sunflecks, and canopy architecture in mesic shrub thickets

Expansion of shrubs into grasslands is often accompanied by a reduction in understory light and an associated reduction of shade-intolerant species. However, effects of specific canopy architectural characteristics on the light environment under shrub thickets are unknown. Our objective was to determine what characteristics of canopy architecture most influence understory light in monospecific shrub thickets. We quantified understory light and canopy architecture for five shrub species in the eastern United States that have a history of expansion, and we used multiple regression to determine which canopy characteristics best predicted light attenuation and relative contribution of sunflecks. Measurements included leaf angle, leaf azimuth, branch bifurcation ratio, leaf area index (LAI), canopy depth (the vertical distance from the bottommost leaf to the top of the canopy), and leaf area density (LAD) as well as understory photosynthetic photon flux density (PPFD). The best predictor of light attenuation and the occurrence of sunflecks for all species was canopy depth. Multiple leaf and plant-level traits were correlated with canopy depth but not with LAI or LAD. The invasive shrub Elaeagnus umbellata had the lowest understory light levels of the species examined although LAI values for Morella cerifera and Rhododendron maximum were higher. Branch bifurcation ratios for E. umbellata were significantly higher than for other species and this likely contributed to the differences in light attenuation and suppression of sunflecks. The potential of shrubs to intercept light is primarily dependent on vertical distribution of leaves in the canopy which is itself correlated with fine-scale, species-specific variations in leaf display.     

The role of nitrogen in a simple scheme to scale up photosynthesis from leaf to canopy

A simple analytical scheme, involving the distribution of nitrogen, to scale up photosynthesis from leaf to canopy is proposed. The scheme is based on the assumption that there are two pools of nitrogen in leaves: nitrogen in photosynthetic, degradable structures (N_p) and nitrogen in non-photosynthetic and non-degradable structures (N_s). The rate of photon-saturated photosynthesis, F_m, is assumed to be proportional to N_p and is distributed inside the canopy similarly to photon flux density (PFD). Prior assumptions of an optimum distribution of nitrogen are not a prerequisite. Calculations made with the scheme lead to development of the hypothesis that the canopy can be treated as a ‘big leaf’ on the time scales involved in acclimation of photosynthesis to PFD. Simulations using parameters for tree species with different requirements for PFD show that shade-tolerant species may have denser canopies than sun-demanding species because of smaller amounts of non-photosynthetic structural nitrogen and/or supporting tissue in their leaves.     

Estimating stand volume in broad-leaved forest using discrete-return LiDAR: plot-based approach

Quantitative assessment of forests is important at a variety of scales for forest planning and management. This study investigated the use of small-footprint discrete-return lidar for estimating stand volume in broad-leaved forest at plot level. Field measurements were conducted at 20 sample plots in the study area in western Japan, composed of temperate broad-leaved trees. Five height variables and two density variables were derived from the lidar data: 25th, 50th, 75th, and 100th percentiles, and mean of laser canopy heights as height variables (h ₂₅, h ₅₀, h ₇₅, h ₁₀₀, h _mean); and ground fraction and only-and-vegetation fraction (d _GF, d _OVF) as density variables, defined respectively as the proportion of laser returns that reached the ground, and the proportion of only echoes (i.e., single pulse returns for which the first and last pulses returned from the same point) within vegetation points. In addition, the normalized difference vegetation index (NDVI), which is often used as an estimator for leaf area index (LAI) and above-ground biomass, was derived from multispectral digital imagery as an alternative density variable (d _NDVI). Nonlinear least-square regression with cross-validation analysis was performed with single variables and combinations; a total of 23 models were studied. The best prediction was found when h ₇₅ and d _OVF were used as independent variables, resulting in adjusted R ² of 0.755 and root-mean-square error (RMSE) of 41.90 m³ ha⁻¹, corresponding to 16.4% of the mean stand volume, better than or comparable to the prediction models of previous studies.     

3D lidar imaging for detecting and understanding plant responses and canopy structure

Understanding and diagnosing plant responses to stress will benefit greatly from three-dimensional (3D) measurement and analysis of plant properties because plant responses are strongly related to their 3D structures. Light detection and ranging (lidar) has recently emerged as a powerful tool for direct 3D measurement of plant structure. Here the use of 3D lidar imaging to estimate plant properties such as canopy height, canopy structure, carbon stock, and species is demonstrated, and plant growth and shape responses are assessed by reviewing the development of lidar systems and their applications from the leaf level to canopy remote sensing. In addition, the recent creation of accurate 3D lidar images combined with natural colour, chlorophyll fluorescence, photochemical reflectance index, and leaf temperature images is demonstrated, thereby providing information on responses of pigments, photosynthesis, transpiration, stomatal opening, and shape to environmental stresses; these data can be integrated with 3D images of the plants using computer graphics techniques. Future lidar applications that provide more accurate dynamic estimation of various plant properties should improve our understanding of plant responses to stress and of interactions between plants and their environment. Moreover, combining 3D lidar with other passive and active imaging techniques will potentially improve the accuracy of airborne and satellite remote sensing, and make it possible to analyse 3D information on ecophysiological responses and levels of various substances in agricultural and ecological applications and in observations of the global biosphere.     

Chronosequence predictions are robust in a Neotropical secondary forest,but plots miss the mark

Tropical secondary forests (TSF) are a global carbon sink of 1.6 Pg C/year. However, TSF carbon uptake is estimated using chronosequence studies that assume differently aged forests can be used to predict change in aboveground biomass density (AGBD) over time. We tested this assumption using two airborne lidar datasets separated by 11.5 years over a Neotropical landscape. Using data from 1998, we predicted canopy height and AGBD within 1.1 and 10.3% of observations in 2009, with higher accuracy for forest height than AGBD and for older TSFs in comparison to younger ones. This result indicates that the space‐for‐time assumption is robust at the landscape‐scale. However, since lidar measurements of secondary tropical forest are rare, we used the 1998 lidar dataset to test how well plot‐based studies quantify the mean TSF height and biomass in a landscape. We found that the sample area required to produce estimates of height or AGBD close to the landscape mean is larger than the typical area sampled in secondary forest chronosequence studies. For example, estimating AGBD within 10% of the landscape mean requires more than thirty 0.1 ha plots per age class, and more total area for larger plots. We conclude that under‐sampling in ground‐based studies may introduce error into estimations of the TSF carbon sink, and that this error can be reduced by more extensive use of lidar measurements.     

Characterization of the canopy structure of Dutch grasslands

In order to characterize the canopy structure of different grassland types 50 stands, representing 15 syntaxonomically distinct types, were examined by harvesting the standing crop during the main flowering period. The types differ in water and nutrient conditions and vary largely in aboveground phytomass (0.5–23 t·ha^?1) and Leaf Area Index (0.4–21: bifacial). Living aboveground phytomass and canopy structure are almost entirely determined by phanerogams. The graminoids generally dominate over the forbs. Variation in aboveground phytomass is related to foliage characteristics such as Aboveground Leaf Area Ratio, Specific Foliage Weight and Leaf Area Development. A PCA with these canopy variables and the variables canopy height, phytomass density and the phytomass ratios stem-leaf-inflorescence shows a clear arrangement related to aboveground phytomass and LAI. The grassland stands can be divided into four groups of different productivity levels for which various combinations of canopy variables are characteristic. Leaf size and leaf inclination are also used for characterization of the different grasslands. Ordinations with these variables by means of PCA and CCA resemble partly to those computed with the canopy variables of aboveground phytomass and LAI. Small leaf sizes are characteristic for low productive grasslands, while the largest leaves occur in high productive grasslands, although they mostly do not belong to the species contributing most strongly to the phytomass of the stand. The leaf inclinations erect and erecto-patent are most common in each grassland. Horizontal leaf areas occur less frequent, but they are relatively well presented in some high productive grasslands. Ranges in leaf size vary more than ranges in leaf inclination do for this series of grasslands. However, leaf size and leaf inclination are useful variables for characterization of grassland canopies in a hierarchical way, where phytomass and leaf area are the first criteria for such a characterization.     

Canopy photosynthetic production in a Japanese larch forest. II. Estimation of the canopy photosynthetic production

Seasonal changes and yearly gross canopy photosynthetic production were estimated for an 18 year old Japanese larch (Larix leptolepis) forest between 1982 and 1984. A canopy photosynthesis model was applied for the estimation, which took into account the effect of light interception by the non-photosynthetic organs. Seasonal changes in photosynthetic ability, amount of canopy leaf area and light environment within the canopy were also taken into account. Amount of leaf area was estimated by the leaf area growth of a single leaf. The change of light environment within the canopy during the growing season was estimated with a light penetration model and the leaf increment within the canopy. Canopy respiration and surplus production were calculated as seasonal and yearly values for the three years studied. Mean yearly estimates of canopy photosynthesis, canopy respiration and surplus production were 37, 13 and 23 tCO₂ ha⁻¹ year⁻¹, respectively. Vertical trend, seasonal changes and yearly values of the estimates were analyzed in relation to environmental and stand factors.     

基于PROSAIL辐射传输模型的毛竹林分冠层反射率模拟研究

冠层光谱反射率直接关系到毛竹（Phyllostachys pubescens Mazel）林冠层参数的反演,对毛竹林地土壤肥力间接估测具有重要意义。以PROSPECT5、PROSAIL模型为基础,从叶片尺度和冠层尺度分析模型参数对叶片和冠层反射率的影响,构建毛竹冠层叶面积指数（LAI）-冠层反射率查找表并通过代价函数选取最优冠层反射率,从而实现毛竹林分冠层反射率的准确模拟。结果表明,在叶片尺度,PROSPECT模型参数敏感性从高到低依次为叶肉结构参数（N） > 叶绿素含量（C_ab） > 等效水厚度（EWT） > 干物质含量（C_m） > 类胡萝卜素含量（C_ar）;在冠层尺度,PROSAIL模型参数敏感性从高到低依次为LAI > C_ab > EWT > C_m > N > C_ar > ALA（平均叶倾角）;叶片尺度反射率整体大于冠层尺度反射率;在400~900 nm波长范围内,PROSAIL模型模拟的冠层光谱反射率与实测光谱反射率拟合效果较好,相对误差为6.71%。     

Effect of the orientation of dipole fragments of the phospholipid molecule on the bilayer structure

By the method of minimization of phospholipid bilayers free energy it was shown that at a decrease of inclination angle of dipole fragments of phospholipid molecules to the bilayer surface there take place: a) an increase of bilayer density, b) a decrease of bilayer thickness caused by a decrease of dipole repulsive forces, c) an increase of the number of molecules in the bilayer due to hydrophobization of the latter. The existence of upper and lower critical angles of dipoles inclination to the bilayer surface were found: theta upcrit, theta lowcrit, if theta greater than or equal to the layers must be destroyed, and if theta less than or equal to theta lowcrit, transition of bilayers to the crystalline state must take place.     

The slow reversibility of photosystem II thermal energy dissipation on transfer from high to low light may cause large losses in carbon gain by crop canopies: a theoretical analysis

Regulated thermal dissipation of absorbed light energy within the photosystem II antenna system helps protect photosystem II from damage in excess light. This reversible photoprotective process decreases the maximum quantum yield of photosystem II (Fv)/Fm) and CO2 assimilation (phiCO2), and decreases the convexity of the non-rectangular hyperbola describing the response of leaf CO2 assimilation to photon flux (theta). At high light, a decrease in phiCO2 has minimal impact on carbon gain, while high thermal energy dissipation protects PSII against oxidative damage. Light in leaf canopies in the field is continually fluctuating and a finite period of time is required for recovery of phiCO2 and when light drops below excess levels. Low phiCO2) and can limit the rate of photosynthetic carbon assimilation on transfer to low light, an effect prolonged by low temperature. What is the cost of this delayed reversal of thermal energy dissipation and phiCO2 recovery to potential CO2 uptake by a canopy in the field? To address this question a reverse ray-tracing algorithm for predicting the light dynamics of 120 randomly selected individual points in a model canopy was used to describe the discontinuity and heterogeneity of light flux within the canopy. Because photoprotection is at the level of the cell, not the leaf, light was simulated for small points of 10(4) micro m rather than as an average for a leaf. The predicted light dynamics were combined with empirical equations simulating the dynamics of the light-dependent decrease and recovery of phiCO2 and and their effects on the integrated daily canopy carbon uptake (A'c). The simulation was for a model canopy of leaf area index 3 with random inclination and orientation of foliage, on a clear sky day (latitude 44 degrees N, 120th day of the year). The delay in recovery of photoprotection was predicted to decrease A'c by 17% at 30 degrees C and 32% at 10 degrees C for a chilling-susceptible species, and by 12.8% at 30 degrees C and 24% at 10 degrees C for a chilling-tolerant species. These predictions suggest that the selection, or engineering, of genotypes capable of more rapid recovery from the photoprotected state would substantially increase carbon uptake by crop canopies in the field.