水稻高光谱变化特征与叶绿素含量监测研究

叶绿素含量是评价水稻光合效率的重要指标,实时无损监测叶绿素含量对水稻生长诊断具有重要意义。以水稻P88S(绿叶)和黄1S(黄叶)为试验材料,分析高光谱指数与水稻叶绿素含量的关系,并构建冠层反射光谱与水稻叶绿素含量监测模型。研究结果表明:水稻不同叶色的冠层光谱反射率随着植株生长而不断变化,在绿叶材料P88S的502-711 nm和黄叶材料黄1S的487-716 nm可见光波长范围内,叶绿素含量与一阶微分光谱的相关系数呈极显著正相关。以P88S RVI(363,675)和黄1S DVI'(639,680)作为光谱参数,与叶绿素含量建立估算模型拟合效果最佳,说明利用高光谱技术结合一阶微分光谱的方法可以监测水稻叶绿素含量。     

Anthocyanin contribution to chlorophyll meter readings and its correction

Leaf chlorophyll content is an important physiological parameter which can serve as an indicator of nutritional status, plant stress or senescence. Signals proportional to the chlorophyll content can be measured non-destructively with instruments detecting leaf transmittance (e.g., SPAD-502) or reflectance (e.g., showing normalized differential vegetation index, NDVI) in red and near infrared spectral regions. The measurements are based on the assumption that only chlorophylls absorb in the examined red regions. However, there is a question whether accumulation of other pigments (e.g., anthocyanins) could in some cases affect the chlorophyll meter readings. To answer this question, we cultivated tomato plants (Solanum lycopersicum L.) for a long time under low light conditions and then exposed them for several weeks (4 h a day) to high sunlight containing the UV-A spectral region. The senescent leaves of these plants evolved a high relative content of anthocyanins and visually revealed a distinct blue color. The SPAD and NDVI data were collected and the spectra of diffusive transmittance and reflectance of the leaves were measured using an integration sphere. The content of anthocyanins and chlorophylls was measured analytically. Our results show that SPAD and NDVI measurement can be significantly affected by the accumulated anthocyanins in the leaves with relatively high anthocyanin content. To describe theoretically this effect of anthocyanins, concepts of a specific absorbance and a leaf spectral polarity were developed. Corrective procedures of the chlorophyll meter readings for the anthocyanin contribution are suggested both for the transmittance and reflectance mode.     

基于因子分析的苜蓿叶片叶绿素高光谱反演研究

因子分析是一种能够将具有错综复杂关系的变量归结为少数几个综合因子的多变量统计分析方法,在降低数据维数的同时又可以保存足够的信息,这为处理信息量丰富但冗余较大的高光谱数据提供了一种有效方法。本文利用2010年9月23日采集的16个样点的苜蓿叶片反射率及叶绿素含量数据,采用因子分析方法,分别提取苜蓿叶片反射率光谱400～900nm,以及可见光400nm～760nm和近红外760～900nm光谱区的公共因子,分析因子载荷分布、载荷总量对公共因子与叶绿素含量相关性的影响。利用逐步回归法建立基于公共因子的叶片叶绿素反演模型,并将反演模型与光谱指数建立的模型进行对比。研究表明,1）公共因子与叶片叶绿素的相关性,在更大程度上是与该因子在各个波段上载荷分布有关,而不是总载荷量;2）对波谱进行分区建立的反演模型略优于全区因子分析建立的反演模型;3）与常用于叶片叶绿素含量反演的光谱指数CARI、MCARI、mND680、mND705、mSR705、TVI、DmSR、BGI、BRI相比,因子分析建立的叶绿素反演模型精度更高。     

Leaf spectral signatures differ in plant species colonizing habitats along a hydrological gradient

Aims We aimed at determining differences in the leaf spectral signatures of plant species groups growing in habitats along the hydrological gradient of an intermittent wetland and to define leaf traits that explain their variability. We want to contribute to the understanding of the causes for plant spectrum variability at leaf and community levels.Methods We measured leaf reflectance spectra (300–887nm) of representative plant species from different habitats and analyzed spectral differences among species groups. To explain leaf spectra variability within a group, we performed detailed analyses of leaf morphological and biochemical traits in selected species.Important findings The reflectance spectra of the different species groups differed most in the green, yellow and red spectral ranges. The reflectance spectra of submerged leaves of hydrophytes with simple structures were explained by their biochemical traits (carotenoids), while for more complex aerial leaves, morphological traits were more important. In submerged and natant leaves of amphiphytes, total mesophyll and spongy tissue thickness were the most important traits, and these explained 44% and 47%, respectively, of the spectrum variability of each plant group. In general, the redundancy analysis biplots show that samples of different plant species colonizing the same habitat form separate clusters and are related to the explanatory variables in different ways. The redundancy analysis biplots of helophytes and wet meadow species show clustering of graminoids and dicots into two distinct groups. Leaf encrustation (prickle hair properties and epidermis thickness) is important for graminoids, while leaf thickness and specific leaf area have more important roles in dicots. Our results show that knowledge of the species composition and leaf traits is necessary to interpret the reflectance spectra of such plant communities.     

桉树叶片光合色素含量高光谱估算模型

色素在植物的生理生态过程中非常重要,利用高光谱数据,揭示光谱反射率上特征波段与光合色素含量间的关系将有助于理解光合色素光谱反射特征的规律,同时为利用高光谱遥感技术快速无损监测植物叶片光合色素提供了技术支持.利用野外采集的桉树叶片样本,在实验室内测定了叶片的高光谱反射率及对应的叶绿素、类胡萝卜素含量.利用光谱分析技术和统计学方法对光谱数据进行处理分析,提取了光谱特征参量,并建立叶绿素、类胡萝卜素含量与光谱特征参量间的估算模型.通过精度检验,研究结果表明以(SDr-SDb)/(SDr+SDb)为变量建立的指数模型估算效果最佳.     

Attenuated total reflectance spectroscopy of plant leaves: a tool for ecological and botanical studies

Attenuated total reflectance (ATR) spectra of plant leaves display complex absorption features related to organic constituents of leaf surfaces. The spectra can be recorded rapidly, both in the field and in the laboratory, without special sample preparation. This paper explores sources of ATR spectral variation in leaves, including compositional, positional and temporal variations. Interspecific variations are also examined, including the use of ATR spectra as a tool for species identification. Positional spectral variations generally reflected the abundance of cutin and the epicuticular wax thickness and composition. For example, leaves exposed to full sunlight commonly showed more prominent cutin- and wax-related absorption features compared with shaded leaves. Adaxial vs. abaxial leaf surfaces displayed spectral variations reflecting differences in trichome abundance and wax composition. Mature vs. young leaves showed changes in absorption band position and intensity related to cutin, polysaccharide, and possibly amorphous silica development on and near the leaf surfaces. Provided that similar samples are compared (e.g. adaxial surfaces of mature, sun-exposed leaves) same-species individuals display practically identical ATR spectra. Using spectral matching procedures to analyze an ATR database containing 117 individuals, including 32 different tree species, 83% of the individuals were correctly identified.     

Relationships between leaf chlorophyll content and spectral reflectance and algorithms for non-destructive chlorophyll assessment in higher plant leaves

Leaf chlorophyll content provides valuable information about physiological status of plants. Reflectance measurement makes it possible to quickly and non-destructively assess, in situ, the chlorophyll content in leaves. Our objective was to investigate the spectral behavior of the relationship between reflectance and chlorophyll content and to develop a technique for non-destructive chlorophyll estimation in leaves with a wide range of pigment content and composition using reflectance in a few broad spectral bands. Spectral reflectance of maple, chestnut, wild vine and beech leaves in a wide range of pigment content and composition was investigated. It was shown that reciprocal reflectance (R lambda)-1 in the spectral range lambda from 520 to 550 nm and 695 to 705 nm related closely to the total chlorophyll content in leaves of all species. Subtraction of near infra-red reciprocal reflectance, (RNIR)-1, from (R lambda)-1 made index [(R lambda)(-1)-(RNIR)-1] linearly proportional to the total chlorophyll content in spectral ranges lambda from 525 to 555 nm and from 695 to 725 nm with coefficient of determination r2 > 0.94. To adjust for differences in leaf structure, the product of the latter index and NIR reflectance [(R lambda)(-1)-(RNIR)-1]*(RNIR) was used; this further increased the accuracy of the chlorophyll estimation in the range lambda from 520 to 585 nm and from 695 to 740 nm. Two independent data sets were used to validate the developed algorithms. The root mean square error of the chlorophyll prediction did not exceed 50 mumol/m2 in leaves with total chlorophyll ranged from 1 to 830 mumol/m2.     

植被生化组分光谱模型抗土壤背景能力分析

应用LOPEX'93(Leaf Optical Properties Experiment)数据,分析了统计回归模型在进行植被叶绿素和水分反演中抗土壤背景影响的能力,模型参数分别使用了:反射率及其变化形式、光谱位置变量、植被指数。在LOPEX'93数据库的植被波谱中分别加入10%-90%的实测土壤光谱信息,得到植被与土壤的混合光谱,并分析混合光谱对植被生化组分的响应。结果表明:应用反射率及其变化形式进行植被叶绿素反演时,以730nm和400nm组合的反射率和反射率倒数的对数为参数的模型具有最高的抗土壤背景能力,在土壤背景所占比例从低到高的变化过程中,以二者反射率组合为参数的模型与叶绿素的相关系数,始终保持在0.645附近,以二者反射率倒数的对数为参数的模型与植被叶绿素的相关系数保持在0.650附近;应用反射率及其变化形式进行植被含水量反演时,以1100,1170,1000,1040,1080nm组合的反射率为参数的模型以及以1170,960,1210,1090,1080,950,1220,1210nm反射率倒数的对数组合为参数的模型具有较高的稳定性,在土壤组分变化的过程中,以上模型与植被含水量的相关系数均稳定的高于0.99;对于光谱位置变量的分析中,以红边-绿峰-红谷组合的模型与植被叶绿素含量具有较高、而且稳定的相关系数,在土壤背景所占比例变化的情况下,相关系数稳定在0.53附近;在应用植被指数进行叶绿素的反演过程中,植被指数与叶绿素的相关系数在土壤背景所占比例变化的情况下变化较大,抗土壤背景的能力均较差;在应用植被指数进行植被水分含量的反演时,以水分指数Ratio975和Ratio1200相关系数最高,且在不同比例土壤背景变化下稳定,相关系数分别分布在0.980附近和0.960附近。该结果可用于指导不同植被覆盖条件下植被冠层参数的反演,提高反演的稳定性和准确性。     

Phosphorescence of protochlorophyll(ide) and chlorophyll(ide) in etiolated and greening bean leaves

The assignment is presented for the principal phosphorescence bands of protochlorophyll(ide), chlorophyllide and chlorophyll in etiolated and greening bean leaves measured at -196°C using a mechanical phosphoroscope. Protochlorophyll(ide) phosophorescence spectra in etiolated leaves consist of three bands with maxima at 870, 920 and 970 nm. Excitation spectra show that the 870 nm band belongs to the short wavelength protochlorophyll(ide), P627. The latter two bands correspond to the protochlorophyll(ide) forms, P637 and P650. The overall quantum yield for P650 phosphorescence in etiolated leaves is near to that in solutions of monomeric protochlorophyll, indicating a rather high efficiency of the protochlorophyll(ide) triplet state formation in frozen plant material. Short-term (2–20 min) illumination of etiolated leaves at the temperature range from -30 to 20°C leads to the appearance of new phosphorescence bands at about 990–1000 and 940 nm. Judging from excitation and emission spectra, the former band belongs to aggregated chlorophyllide, the latter one, to monomeric chlorophyll or chlorophyllide. This indicates that both monomeric and aggregated pigments are formed at this stage of leaf greening. After preillumination for 1 h at room temperature, chlorophyll phosphorescence predominates. The spectral maximum of this phosphorescence is at 955–960 nm, the lifetime is about 2 ms, and the maximum of the excitation spectrum lies at 668 nm. Further greening leads to a sharp drop of the chlorophyll phosphorescence intensity and to a shift of the phosphorescence maximum to 980 nm, while the phosphorescence lifetime and a maximum of the phosphorescence excitation spectrum remains unaltered. The data suggest that chlorophyll phosphorescence belongs to the short wavelength, newly synthesized chlorophyll, not bound to chloroplast carotenoids. Thus, the phosphorescence measurement can be efficiently used to study newly formed chlorophyll and its precursors in etiolated and greening leaves and to address various problems arising in the analysis of chlorophyll biosynthesis.Abbreviations Pchl protochlorophyll and protochlorophyllide - Chld chlorophyllide - Chl chlorophyll     

叶绿素a和类胡萝卜素仿生膜研究

在植物叶片的叶绿体中的俘光色素主要有叶绿素和类胡萝卜素。有机溶剂中的叶绿素。很快会发生光解,但即使在夏日炎炎烈日暴晒下,叶片中的叶绿素a却没有任何变化。在叶绿体中叶绿素a和类胡萝卜素的分子构象是抵御强光照的主要原因。本文仿照叶绿体膜结构进行离体模拟实验,采用荧光光谱法来研究强光下β-car保护chla的物理机制。仿照叶绿体膜原理,可以开发出新一代彷生防晒霜。     

Estimation of the leaf chlorophyll content using multiangular spectral reflectance factor

Chlorophyll is one of the primary pigments of plant leaves, and changes in its content can be used to characterize the physiological status of plants. Spectral indices have been devised and validated for estimating leaf chlorophyll content (LCC). However, most of the existing spectral indices do not consider the influence of angular reflection on the accuracy of the LCC estimation. In this study, the spectral reflectance factors of leaves from three plant species were measured from several observations in the principal plane. The relationship between the existing spectral indices and the LCC from different directions suggests that the directional reflection of a leaf surface impacts the accuracy of its LCC estimation. Subsequently, the ratio of reflectance differences, that is, the modified Datt index, was tested to reduce the directional reflection effect when predicting LCC. Our results indicated that the modified Datt index not only estimated LCC with high accuracy for all observation directions and plant species but also consistently predicted the LCC of each species in individual observation directions. Our method opens the possibility for optical detection of LCC using multiangular spectral reflection, which is convenient for plant science studies focused on the variation in LCC.     

Application of Reflectance Spectroscopy for Analysis of Higher Plant Pigments

Nondestructive techniques developed by the authors for assessment of chlorophylls, carotenoids, and anthocyanins in higher plant leaves and fruits are presented. The spectral features of leaf reflectance in the visible and near infrared regions are briefly considered. For pigment analysis only reflectance values at several specific wavelengths are required. The chlorophyll (Chl) content over a wide range of its changes can be assessed during leaf ontogeny using reflectance near 700 nm and, in the absence of anthocyanins, at 550 nm. The approaches used for elimination of Chl interference in the analysis of carotenoids (reflectance at 520 nm) and anthocyanins (at 550 nm) are described. The suitability of reflectance spectroscopy for estimates of carotenoid/chlorophyll ratios during leaf senescence and fruit ripening is demonstrated. The algorithms developed for pigment analysis are presented, and the conditions of their applicability are considered. Further perspectives for the application of reflectance spectroscopy including remote sensing for estimation of plant pigment content and physiological states are discussed.     

不同灌溉量夏玉米叶绿素含量的高光谱特征及其反演

植物叶绿素含量直接影响其光合作用,并与植物的光谱特征密切相关。以夏玉米为研究对象,采用人工控水方法研究了夏玉米七叶期不同灌溉量下冠层叶绿素含量特征及其与光谱特征之间的关系。结果表明：灌溉量越少,夏玉米叶片叶绿素含量越低,冠层光谱反射率越高,绿峰位置"红移",而红边位置"蓝移"。叶绿素含量与光谱特征参数、植被光谱指数之间存在极显著相关关系,据此建立了冠层叶绿素含量高光谱估算模型,且基于植被指数模型较基于单一光谱特征参数模型模拟效果更好。研究结果可为夏玉米叶绿素含量的快速无损测定以及夏玉米干旱监测提供依据。     

基于光谱指数的喀斯特植物叶片叶绿素含量定量估算

为了探讨适合于喀斯特植物叶片叶绿素含量估算的光谱指数,在总结以往基于光谱指数的植物生化参数估算研究基础上发现,常用光谱指数通常采用差值、比值、归一化以及倒数差值方式来构建。因此,我们通过上述4种光谱指数构建方式对所采集的4种典型喀斯特植物——黄荆(Vitex negundo)、盐麸木(Rhus chinensis)、朴树(Celtis sinensis)和红背山麻杆(Alchornea trewioides)叶片原始光谱反射率及其一阶导数值与同步测定的叶片叶绿素含量进行遍历分析,以期获得最优光谱指数并将其应用于喀斯特植物叶片叶绿素含量定量估算研究。结果表明：(1)常用光谱指数中,改良红边归一化指数(modified red-edge normalized difference vegetation index, mND705)对喀斯特植物叶片叶绿素含量估算效果较好(决定系数为0.45,均方根误差为0.26 mg·g^-1)。(2)虽然荧光比值(fluorescence ratio index, FRI1)和叶绿素吸收面积光谱指数(chlorophyll absorp...     

Leaf optical properties in amphibious plant species are affected by multiple leaf traits

The amphibious plant species Sagittaria sagittifolia and Ranunculus lingua here serve as model systems to study differences in leaf optical properties of different leaf types that develop in aquatic and terrestrial environments. We aimed to determine leaf traits that explain most of the variability in the reflectance and transmittance spectra in the range from 280 to 880 nm. Comparisons of leaves of the same form revealed marked differences in their structures and particularly in the content of total methanol-soluble UV-absorbing compounds. Submerged leaves transmit radiation over the whole range measured, but emerged leaves transmit only at wavelengths from 500 to 650 nm, and above 690 nm. Redundancy analysis shows that biochemical leaf traits, namely the UV-absorbing compounds chlorophyll a and b, together with the specific leaf area (SLA), significantly affect the reflectance spectra, explaining 60% of the spectra variability. Pigment levels negatively affect reflectance, while the effect of SLA is positive. Physical traits like thickness of the palisade mesophyll, SLA, and thickness of the lower and upper epidermis, along with anthocyanin content, explain 62% of the transmittance spectra variability. This study provides new insight into the understanding of data collected for aquatic and semi-aquatic plants based on spectral analyses.     

干旱胁迫下雷竹叶片叶绿素的高光谱响应特征及含量估算

植物叶片的反射光谱特征与叶绿素含量密切相关。以重要的笋用竹种雷竹(Phyllostachys violascens)为研究对象,采用盆栽及控水试验方法研究了2年生雷竹在干旱胁迫条件下冠层叶片反射光谱的响应特征,分析了叶片叶绿素含量与不同波段光谱反射率一阶微分值以及光谱特征参数之间的相关关系,并以雷竹叶绿素含量敏感波段及构建的植被指数与叶绿素含量进行了拟合。结果表明,重度缺水处理后雷竹叶片叶绿素含量显著降低,在可见光区叶片光谱反射率随叶绿素含量的降低而增加,以波长493、639、693、756 nm等处的光谱反射率一阶微分值与叶绿素含量的相关性较高。雷竹叶片叶绿素含量与光谱特征参数如绿峰反射率、红谷反射率、蓝边面积、绿峰面积之间的相关性较高。与已有的植被指数相比基于雷竹叶绿素含量敏感波段修正后的植被指数与叶绿素含量相关性优于原植被指数。基于反射率一阶微分值构建的多元回归方程以及修正的绿色归一化植被指数(m GNDVI)构建的回归方程拟合效果较好,为雷竹叶绿素含量的较优估算方程。研究结果可以为雷竹叶绿素含量的快速无损测定以及季节性干旱条件下雷竹林的科学经营及灾后评估提供依据。     

Cladocera in shallow lakes from the Ecuadorian Andes show little response to recent climate change

Floating-leaved rhizophytes and pleustophytes are the first barrier to Sun’s rays and significantly affect the light regime of the water column. To evaluate these effects on light attenuation, the reflectance and transmittance spectra variability were examined according to the leaf traits within three plant groups: (1) seed plants with green abaxial surfaces; (2) seed plants with red abaxial surfaces; and (3) ferns with trichomes. Specific leaf area (SLA), chlorophyll a and b, and UV-B and UV-A-absorbing substances differed between these three groups. The ‘spectral signatures’ of floating-leaved seed plants are similar to those of terrestrial seed plants, with a peak in the green region and a pronounced ‘red edge’. Ferns transmitted more light along the whole spectrum compared to other species. Most reflectance and transmittance spectra variability of the first group was explained by SLA. In the second group, 36% of the reflectance spectra variability was explained by tissue density and carotenoids, and 48% of the transmittance spectra variability by carotenoids, anthocyanins and SLA. In ferns, the reflectance spectra variability was mainly explained by chlorophylls, and partly by trichome length and mesophyll thickness, with the transmittance spectra variability significantly affected by chlorophyll b.     

Leaf optical properties in higher plants: linking spectral characteristics to stress and chlorophyll concentration

A number of studies have linked responses in leaf spectral reflectance, transmittance, or absorptance to physiological stress. A variety of stressors including dehydration, flooding, freezing, ozone, herbicides, competition, disease, insects, and deficiencies in ectomycorrhizal development and N fertilization have been imposed on species ranging from grasses to conifers and deciduous trees. In all cases, the maximum difference in reflectance within the 400-850 nm wavelength range between control and stressed states occurred as a reflectance increase at wavelengths near 700 nm. In studies that included transmittance and absorptance as well as reflectance, maximum differences occurred as increases and decreases, respectively, near 700 nm. This common optical response to stress could be simulated closely by varying the chlorophyll concentration of model leaves (fiberglass filter pads) and by the natural variability in leaf chlorophyll concentrations in senescent leaves of five species. The optical response to stress near 700 nm, as well as corresponding changes in reflectance that occur in the green-yellow spectrum, can be explained by the general tendency of stress to reduce leaf chlorophyll concentration.     

Functional relationship between chlorophyll content and leaf reflectance, and light-capturing efficiency of Japanese forest species

We examined the functional relationship between chlorophyll concentrations and light spectral absorption in 16 species of woody, vine and herbaceous plants in northern Japan. Leaves of each species from under forest shade and in more open sites were measured for chlorophyll, specific leaf area (SLA) and spectral absorption. In all species, SLA increased and the Chl a : b ratio declined in shade- vs open-grown leaves indicating an adaptive adjustment to forest shade in these leaf characters. However, the expected increase in the ratio of 680 to 700 nm absorption in shade leaves did not occur in all species. Light absorption at 680 relative to 700 nm was lower in the shade leaves of Acer japonicum. Kalopanax pictus, Panax japonicus and Petasites japonicus even with a reduced Chl a : b , a commonly accepted indicator of shade adaptation. Therefore, spectral measurements in these species failed to support Chl concentrations that were expected to confer an improvement in the absorption of red light (<680nm) deficient relative to far-red light (>700 nm) in the forest shade. Compared with other species, the absorption pattern of these four 'non-conforming' species is associated with a higher ratio of shade:open leaves in reflectance spectra in the 600–750 nm range. This suggests an increased reflectance in shade leaves caused by changes in leaf surface properties which are not immediately apparent. We conclude that adaptive spectral absorption cannot always be inferred from changes in specific leaf area and chlorophyll a and b concentrations.     

不同水氮条件下水稻冠层反射光谱与植株含水率的定量关系

研究了不同土壤水氮条件下水稻 (Oryzasativa) 冠层光谱反射特征和植株水分状况的量化关系。结果表明, 水稻冠层近红外光谱反射率随土壤含水量的降低而降低, 短波红外光谱反射率随土壤含水量的降低而升高。相同土壤水分条件下, 高氮水稻的冠层含水率高于低氮水稻的冠层含水率 ;同一水分条件下, 高氮处理的可见光区和短波红外波段光谱反射率低于低氮处理, 近红外波段光谱反射率高于低氮处理。发现拔节后比值植被指数 (R810 /R460 ) 与水稻叶片含水率和植株含水率呈极显著的线性相关, 模型的检验误差 (RootmeansquareError, RMSE) 分别为 0.93和 1.5 0。表明比值植被指数R810 /R460 可以较好地监测不同生育期水稻叶片和植株含水率。