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。     

Estimating foliar biochemistry from hyperspectral data in mixed forest canopy

Estimating canopy biochemical composition in mixed forests at the level of tree species represents a critical tool for a better understanding and modeling of ecosystem functioning since many species exhibit differences in functional attributes or decomposition rates. We used airborne hyperspectral data to estimate the foliar concentration of nitrogen, carbon and water in three mixed forest canopies in Switzerland. With multiple linear regression models, continuum-removed and normalized HyMap spectra were related to foliar biochemistry on an individual tree level. The six spectral wavebands used in the regression models were selected using both an enumerative branch-and-bound (B&B) and a forward search algorithm. The models estimated foliar concentrations with adjusted R² values between 0.47 and 0.63, based on the best-sampled study site. Regression models composed of wavebands selected by the B&B algorithm always performed better than those developed with forward search. When extrapolating nitrogen concentrations from one to another study site, regression models solely based on causal wavebands (known from literature) mostly outperformed models based on all wavebands. The study demonstrates the potential of both the use of causal wavebands and of the B&B algorithm.     

Influence of foliar phenology and shoot inclination on annual photosynthetic gain in individual beech saplings: A functional–structural modeling approach

We developed a functional–structural plant model for Fagus crenata saplings and calculated annual photosynthetic gains to determine the influences of foliar phenology and shoot inclination on the carbon economy of saplings. The model regenerated the three-dimensional shoot structure and spatial and temporal display of leaves; we calculated the hourly light interception of each leaf with a detailed light model that allowed us to estimate hourly leaf photosynthetic gain taking leaf age into account. To evaluate the importance of simultaneous foliar phenology and slanting shoots in beech saplings, we calculated the photosynthetic budgets for saplings with contrasting foliar phenologies and shoot inclinations. In our simulations, we distinguished between simultaneous and successive foliar phenologies, upright and slanting shoot inclinations, and environments with and without a vertical gradient in light intensity. Other model parameters (including photosynthesis vs. light curve, leaf size, and leaf shape) were obtained directly from live beech saplings. With no vertical gradient in light intensity, modeled saplings with simultaneous foliar phenology and slanting shoots (as in live beech) had larger annual photosynthetic gains than saplings with other combinations of traits. Hence, simultaneous foliar phenology and slanting shoots are efficient ways to display leaves in the shaded forest understory light regime where beech saplings thrive. In the presence of vertical light gradients, which can occur in canopy gaps, saplings with upright shoots had larger annual photosynthetic gains than counterparts with slanting shoots. Although mean daily photosynthetic gains of saplings with successive foliar phenology were elevated by exposing leaves to strong light when young and productive, the annual photosynthetic budget of these saplings was reduced (compared to saplings with simultaneous foliar phenology) by their relatively short leaf lifespan. Overall, our results suggest that slanting shoots with simultaneous foliar phenology are particularly successful in shaded environments, where beech often dominates, because they appear to maximize the annual carbon budget by avoiding self-shading and extending leaf lifespans.     

Assessment of allometric algorithms for estimating leaf biomass, leaf area index and litter fall in different-aged Sitka spruce forests

The relationship between leaf area and diameter at breast height(d.b.h.) or sapwood area (AS) has been used to estimate standleaf area or biomass of forest canopies. It has been suggestedthat intra-specific variations in the relationship between standleaf area and d.b.h. or AS can introduce a systematic errorin these estimates for younger and older stands unless additionalparameters relating to canopy structure are included in allometricfunctions. We collected data from a Sitka spruce chronosequenceto parametrize and test different algorithms for the estimationof foliar biomass (FB) and litter inputs over a range of forestages. FB estimates were significantly improved when additionalbiometric information relating to crown structure (canopy opennessand height of live crown) was included in the models. Althoughthe use of the relationship between leaf area and AS for theestimation of leaf area is justified by theoretical considerations(pipe model theory), we show that d.b.h. and other canopy parametersprovided the most robust estimation of leaf area across different-agedstands. Our results also suggest that the accuracy of litterinput estimates depends on needle retention time and annualturnover rate, particularly immediately before and after canopyclosure.     

Seasonal patterns of cytokinins and microclimate and the mediation of gas exchange among canopy layers of mature Acer saccharum trees

Seasonal patterns of cytokinins (CKs) and microclimate were examined in the upper, middle and lower canopy layers of mature Acer saccharum Marsh. (sugar maple) trees to elucidate the potential role of CKs in the mediation of gas exchange. The upper canopy showed a distinctly dissimilar microclimate from the middle and lower canopy layers with higher photosynthetically active radiation and wind speed, but showed no corresponding differences in transpiration (E) or stomatal conductance (g(s)). Although E and g(s) tended to be higher in the upper canopy than in the middle and lower canopies, the differences were not significant, indicating regulation beyond the passive response to changes in microclimate. The upper canopy accumulated significantly higher concentrations of CKs, predominantly as ribosides, and all canopy layers showed distinct seasonal patterns in CK profiles. Multiple regression models showed significant relationships between both g(s) and E and foliar CK concentration, although these relationships varied among canopy layers. The relationships were strongest in the middle and lower canopy layers where there was less fluctuation in leaf water status and less variability in abiotic variables. The relationships between gas exchange parameters and leaf CK concentration began to decouple near the end of the growing season as foliar phytohormone concentrations changed with the approach of dormancy.     

Genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition and tree growth with clones of the F1 hybrid between slash pine and Caribbean pine

The objectives of this study were: (1) to quantify the genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition (δ¹³C) and tree growth of 122 clones of ca. 4-year-old F₁ hybrids between slash pine (Pinus elliottii Engelm var. elliottii) and Caribbean pine (P. caribaea var. hondurensis Barr. et Golf.) grown at two experimental sites with different water and nutrient availability in southeast Queensland, Australia and (2) to examine the potential of using foliar nutrient concentration of the 4-year-old tree canopies for selecting elite F₁ hybrid pine clones with improved nutrient-use efficiency (NUE) and water-use efficiency (WUE), and ultimately enhanced tree growth under ambient growing conditions. There were significant differences in foliar nutrient concentrations between two canopy positions (upper outer and lower outer canopy) sampled, between summer and winter, and between the two sites. This highlights that foliar nutrient concentrations are influenced by sampling and environment. Significant genetic variations in foliar nutrient concentrations were detected between the clones, between the female parents, and between the male parents of the clones in both sampling seasons at both sites. Depending on the nutrient concerned, canopy position, season, and site sampled, the clones accounted for 4.7–33.9% of the total variation in foliar nutrient concentrations, the clone female parents for 0–25.1% and the clone male parents for 0–28.6%. The site-by-clone interactions were statistically significant for foliar N, P, Mg, Cu, Zn, Mn, Fe and mineral concentrations at the upper outer canopy in summer, and for foliar N concentration in winter. There were significant, positive correlations between clone means of foliar δ¹³C and N concentration at the upper outer canopy in summer for the wet site, while clone foliar δ¹³C was also positively related to clone foliar N concentration at both canopy positions in summer for the dry site. This suggests that clone WUE as reflected in foliar δ¹³C may be improved by selecting elite clones with higher foliar N concentration and increased photosynthesis, leading to enhanced tree growth when both water and N are the major growth-limiting factors. This is supported by the positive correlation detected between clone tree height and foliar N concentration at the upper outer canopy for both sites. Thus, foliar nutrient (particularly N) concentration, together with foliar δ¹³C, may be useful for assisting in selection of exotic pine clones with improved NUE and WUE, and enhanced tree growth under the nutrient- and water-limiting environments.     

Interspecific and environmentally induced variation in foliar dark respiration among eighteen southeastern deciduous tree species

We measured variations in leaf dark respiration rate (Rd) and leaf nitrogen (N) across species, canopy light environment, and elevation for 18 co-occurring deciduous hardwood species in the southern Appalachian mountains of western North Carolina. Our overall objective was to estimate leaf respiration rates under typical conditions and to determine how they varied within and among species. Mean dark respiration rate at 20 degrees C (Rd,mass, micromol CO2 per kg leaf dry mass per s) for all 18 species was 7.31 micromol per kg per s. Mean Rd,mass of individual species varied from 5.17 micromol per kg per s for Quercus coccinea Muenchh. to 8.25 micromol per kg per s for Liriodendron tulipifera L. Dark respiration rate varied by leaf canopy position and was higher in leaves collected from high-light environments. When expressed on an area basis, dark respiration rate (Rd,area, micromol CO2 per kg leaf dry area per s) showed a strong linear relationship with the predictor variables leaf nitrogen (Narea, g N per square m leaf area) and leaf structure (LMA, g leaf dry mass per square m leaf area) (r squared = 0.62). This covariance was largely a result of changes in leaf structure with canopy position; smaller thicker leaves occur at upper canopy positions in high-light environments. Mass-based expression of leaf nitrogen and dark respiration rate showed that nitrogen concentration (Nmass, mg N per g leaf dry mass) was only moderately predictive of variation in Rd,mass for all leaves pooled (r squared = 0.11), within species, or among species. We found distinct elevational trends, with both Rd,mass and Nmass higher in trees originating from high-elevation, cooler growth environments. Consideration of interspecies differences, vertical gradients in canopy light environment, and elevation, may improve our ability to scale leaf respiration to the canopy in forest process models.     

森林郁闭度定量估测遥感比值波段的选择

对TM遥感图像各原始及常用比值波段的信息量大小、相互间存在的多重相关性及其对郁闭度估测造成的不利影响进行分析研究。针对特定监测区域,对设置的遥感比值波段,采用平均残差平方和准则进行筛选,最终确定对郁闭度估测有一定影响的遥感波段,并形成可操作的软件系统。     

Variation in morphological and biochemical O3 injury attributes of mature Jeffrey pine within canopies and between microsites

Crown morphology and leaf tissue chemical and biochemical attributes associated with ozone (O3) injury were assessed in the lower, mid- and upper canopy of Jeffrey pine (Pinus jeffreyi Grev. & Balf.) growing in mesic and xeric microsites in Sequoia National Park, California. Microsites were designated mesic or xeric based on topography and bole growth in response to years of above-average precipitation. In mesic microsites, canopy response to O3 was characterized by thinner branches, earlier needle fall, less chlorotic leaf mottling, and lower foliar antioxidant capacity, especially of the aqueous fraction. In xeric microsites, canopy response to O3 was characterized by higher chlorotic leaf mottling, shorter needles, lower needle chlorophyll concentration, and greater foliar antioxidant capacity. Increased leaf chlorotic mottle in xeric microsites was related to drought stress and increased concurrent internal production of highly reactive oxygen species, and not necessarily to stomatal O3 uptake. Within-canopy position also influenced the expression of O3 injury in Jeffrey pine.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Foliar delta(13)C and delta(18)O reveal differential physiological responses of canopy foliage to pre-planting weed control in a young spotted gum (Corymbia citriodora subsp. Variegata) plantation

Weed control may improve the growth of forest plantations by influencing soil water and nutrient availability, but our knowledge of leaf-level physiological responses to weed control at different within-canopy positions is limited for tropical and subtropical plantations. Foliar carbon (delta(13)C) and oxygen (delta(18)O) isotope compositions, gas exchange, and nitrogen (N(mass)) and phosphorus (P(mass)) concentrations at four canopy positions were assessed in a young spotted gum (Corymbia citriodora subsp. Variegata (F. Muell.) A.R. Bean & M.W. McDonald) plantation subjected to either weed control or no weed control treatment, to test if leaves at different positions within the tree canopy had the same physiological responses to the weed control treatment. Weed control increased foliar delta(13)C but lowered delta(18)O in the upper-outer and upper-inner canopy, indicating that weed control resulted in a higher foliar photosynthetic capacity at upper-canopy positions, a conclusion confirmed by gas exchange measurements. The increased photosynthetic capacity resulting from weed control can be explained by an increase in foliar N(mass). In the lower-outer canopy, weed control reduced foliar delta(13)C while lowering delta(18)O even more than in the upper-canopy, suggesting strong enhancement of the partial pressure of CO(2) in the leaf intercellular spaces and of foliar stomatal conductance in lower-canopy foliage. This conclusion was supported by gas exchange measurements. Foliar photosynthesis in the lower-inner canopy showed no significant response to weed control. The finding that leaves at different canopy positions differ in their physiological responses to weed control highlights the need to consider the canopy position effect when examining competition for soil nutrient and water resources between weeds and trees.     

Shoot biomass growth is related to the vertical leaf nitrogen gradient in Salix canopies

Plant canopy optimization models predict that leaf nitrogen (N) distribution in the canopy will parallel the vertical light gradient, and numerous studies with many species have confirmed this prediction. Further, it is predicted that for a given canopy leaf area, a low vertical light extinction coefficient will promote rapid growth. Therefore, the ideal canopy of fast-growing plants should combine high leaf area index with a low light extinction coefficient; the latter being reflected in a flat vertical leaf N gradient throughout the canopy. Based on data from an experimental Salix stand (six varieties) grown on agricultural land in central Sweden, we tested the hypothesis that shoot growth is correlated with vertical leaf N gradient in canopies of hybrid willows bred for biomass production, which could have implications for Salix breeding. Tree improvement research requires screening of growth-related traits in large numbers of plants, but assessment of canopy leaf N gradients by chemical analysis is expensive, time-consuming and destructive. An alternative to analytical methods is to estimate leaf N gradients nondestructively with an optical chlorophyll meter (SPAD method). Here we provide a specific calibration for interpreting SPAD data measured in hybrid willows grown in biomass plantations on fertile agricultural land. Based on SPAD measurements, a significant and inverse relationship (r(2) = 0.88) was found between shoot biomass growth and vertical leaf N gradient across canopies of six Salix varieties.     

Branch growth and gas exchange in 13-year-old loblolly pine (Pinus taeda) trees in response to elevated carbon dioxide concentration and fertilization

We used whole-tree, open-top chambers to expose 13-year-old loblolly pine (Pinus taeda L.) trees, growing in soil with high or low nutrient availability, to either ambient or elevated (ambient + 200 micromol mol-1) carbon dioxide concentration ([CO2]) for 28 months. Branch growth and morphology, foliar chemistry and gas exchange characteristics were measured periodically in the upper, middle and lower crown during the 2 years of exposure. Fertilization and elevated [CO2] increased branch leaf area by 38 and 13%, respectively, and the combined effects were additive. Fertilization and elevated [CO2] differentially altered needle lengths, number of fascicles and flush length such that flush density (leaf area/flush length) increased with improved nutrition but decreased in response to elevated [CO2]. These results suggest that changes in nitrogen availability and atmospheric [CO2] may alter canopy structure, resulting in greater foliage retention and deeper crowns in loblolly pine forests. Fertilization increased foliar nitrogen concentration (N(M)), but had no consistent effect on foliar leaf mass (W(A)) or light-saturated net photosynthesis (A(sat)). However, the correlation between A(sat) and leaf nitrogen per unit area (N(A) = W(A)N(M)) ranged from strong to weak depending on the time of year, possibly reflecting seasonal shifts in the form and pools of leaf nitrogen. Elevated [CO2] had no effect on W(A), N(M) or N(A), but increased A(sat) on average by 82%. Elevated [CO2] also increased photosynthetic quantum efficiency and lowered the light compensation point, but had no effect on the photosynthetic response to intercellular [CO2], hence there was no acclimation to elevated [CO2]. Daily photosynthetic photon flux density at the upper, middle and lower canopy position was 60, 54 and 33%, respectively, of full sun incident to the top of the canopy. Despite the relatively high light penetration, W(A), N(A), A(sat) and R(d) decreased with crown depth. Although growth enhancement in response to elevated [CO2] was dependent on fertilization, [CO2] by fertilization interactions and treatment by canopy position interactions generally had little effect on the physiological parameters measured.     

卫星遥感技术在森林病虫害监测中的应用

本文阐述了利用先进的卫星遥感技术手段，替代传统目视地面调查方法的必要性和可行性。回顾并总结了近２０年来卫星遥感技术在森林病虫害监测领域的研究进展。展望了卫星遥感监测森林病虫害的前景。     

结合Landsat 8与PALSAR-2影像的龙南县针叶林蓄积量遥感估测研究

林分蓄积量估测是林业遥感的重要研究领域,由于云雾天气和光谱饱和现象等因素限制了光学遥感影像估测林分蓄积量的精度。合成孔径雷达(SAR)具有穿透性强、受云雾影响小等特点,弥补了光学遥感的不足。以江西省龙南县的针叶林为研究对象,结合Landsat 8与PALSAR-2双极化SAR影像数据,在遥感数据预处理基础上,提取了光谱信息、植被指数、纹理信息和后向散射系数等共245个遥感因子。基于Pearson相关系数法和多元逐步回归法,筛选出65个遥感因子参与林分蓄积量估测。以林分郁闭度作为分层因子,分别采用线性、KNN、支持向量机(SVM)、多重感知机(MLP)和随机森林(RF)5种模型估测林分蓄积量,并对估测结果进行精度检验。实验结果表明:1)相比单独使用Landsat 8的光谱和纹理信息,基于郁闭度分级并融合PALSAR-2的后向散射信息明显提高了蓄积量的反演精度;2)对于低郁闭度林分,线性模型精度最高(rRMSE=21.16%),中郁闭度林分,多重感知机模型估测效果最好(rRMSE=30.61%),高郁闭度林分,多重感知机模型估测效果最好(rRMSE=27.53%)。在结合PALSAR-2的后向散射系数的基础上,郁闭度分层能有效改善中高蓄积量区域的反演精度。     

红树植物地面反射光谱特征研究

结合使用野外便携式光谱仪对九龙江口红树林保护区白骨壤、秋茄、桐花树3种红树植物的叶冠表观光谱的测量结果,系统分析比较了红树植物叶冠反射光谱特征及其差异;并在一个潮间带底质遥感信息处理平台上对研究地点进行基于表观光谱特征的白骨壤和秋茄的SPOT卫星遥感图像分类;最后讨论了红树光谱研究中存在的问题。     

结合图像纹理特征的森林郁闭度遥感估测

在光谱等传统特征的基础上,结合遥感图像的纹理特征估测郁闭度:首先基于面向地块的方法计算图像的灰度共生矩阵纹理特征,然后用主成分方法分析相关性并降维,最后将图像纹理特征和光谱地形等特征一起作为自变量引入到郁闭度估测的逐步回归模型中。结果表明:结合图像纹理特征的方法比传统的只基于光谱或地形特征的方法在估测精度上有较大提高,判别系数R珔2从0.737提高到0.805,估测精度从81.03%提高到84.32%。     

Chlorophyll content in eucalypt vegetation at the leaf and canopy scales as derived from high resolution spectral data

The physiological status of forest canopy foliage is influenced by a range of factors that affect leaf pigment content and function. Recently, several indices have been developed from remotely sensed data that attempt to provide robust estimates of leaf chlorophyll content. These indices have been developed from either hand-held spectroradiometer spectra or high spectral resolution (or hyperspectral) imagery. We determined if two previously published indices (Datt 1999), which were specifically developed to predict chlorophyll content in eucalypt vegetation by remote sensing at the leaf scale, can be extrapolated accurately to the canopy. We derived the two indices from hand-held spectroradiometer data of eucalypt leaves exhibiting a range of insect damage symptoms. We also derived the indices from spectra obtained from high spectral and spatial resolution Compact Airborne Spectrographic Imager 2 (CASI-2) imagery to determine if reasonable estimates at a scale of < 1 m can be achieved. One of the indices (R 850/R 710 index, where R is reflectance) derived from hand-held spectroradiometer data showed a moderate correlation with relative leaf chlorophyll content (r = 0.59, P < 0.05) for all dominant eucalypt species in the study area. The R (850)/R (710) index derived from CASI-2 imagery yielded slightly lower correlations over the entire data set (r = 0.42, P < 0.05), but correlations for individual species were high (r = 0.77, P < 0.05). A scaling analysis indicated that the R (850)/R (710) index was strongly affected by soil and water cover types when pixels were mixed, but appeared to be invariant to changes in proportions of understory, which may limit its application.     

Mapping dead wood distribution in a temperate hardwood forest using high resolution airborne imagery

Dead wood, in the form of coarse woody debris and standing dead wood, or snags, is an essential structural component of forest ecosystems. It plays a key role in nutrient cycling, ecosystem functions and provision of habitat for a wide variety of species. In order to manage dead wood in a temperate hardwood forest, an understanding of its availability and spatial distribution is important. This research evaluates airborne digital camera remote sensing for mapping temperate forest dead wood across an area within Gatineau Park, Canada. Two approaches were evaluated: (1) direct detection and mapping of canopy dead wood (dead branches and tall snags) through the combination of three techniques in a hybrid classification: ISODATA clustering, object-based classification, and spectral unmixing, and (2) indirect modelling of coarse woody debris and snags using spectral and spatial predictor variables extracted from the imagery. Indirect modelling did not provide useful results while direct detection was successful with field validation showing 94% accuracy for detected canopy level dead wood objects (i.e. 94% of validation sites with canopy dead wood were detected correctly) and 90% accuracy for control sites (i.e. 90% of validation sites with no canopy level dead wood were identified correctly). The procedures presented in this paper are repeatable and could be used to monitor dead wood over time, potentially contributing to applications in forest carbon budget estimation, biodiversity management, and forest inventory.     

落叶松锉叶蜂为害的松林光谱特征分析

通过对不同程度危害的落叶松反射光谱进行测定,研究了不同程度危害的落叶松在绿光区、红光区和近红外区反射光谱的变化特征,并对光谱反射曲线进行微分分析。结果表明,绿光区、红光区和近红外区的落叶松光谱反射率随危害程度的加重分别呈现下降、上升和下降的趋势;对反射率曲线进行微分分析,健康落叶松、轻度、中度和重度为害后的一阶导数光谱反射率最大值随着危害程度增加而下降,并且向短波方向移动(蓝移)。实测光谱数据提取的归一化植被指数(NDVI)与落叶松锉叶蜂危害程度呈显著负相关,对应用遥感技术早期监测落叶松锉叶蜂灾害具有重要指示意义。