RESPONSES OF LEAF SPECTRAL REFLECTANCE TO PLANT STRESS

Leaf spectral reflectances were measured to determine whether leaf reflectance responses to plant stress may differ according to the agent of stress and species. As a result of decreased absorption by pigments, reflectance at visible wavelengths increased consistently in stressed leaves for eight stress agents and among six vascular plant species. Visible reflectance was most sensitive to stress in the 535–640-nm and 685–700-nm wavelength ranges. A sensitivity minimum occurred consistently near 670 nm. Infrared reflectance was comparatively unresponsive to stress, but increased at 1,400–2,500 nm with severe leaf dehydration and the accompanying decreased absorption by water. Thus, visible rather than infrared reflectance was the most reliable indicator of plant stress. Visible reflectance responses to stress were spectrally similar among agents of stress and species.     

冠层水平互花米草叶片光合色素含量的高光谱遥感估算模型

以闽江河口鳝鱼滩湿地互花米草(Spartina alterniflora)的实测冠层高光谱反射率和叶片光合色素含量(LPPC)为数据源,在分析LPPC与原始光谱反射率、一阶导数光谱反射率、22种已报道光谱指数和14种新构建的植被指数相关性的基础上,利用直线回归、指数回归、对数回归以及乘幂回归方法,系统地比较了36种植被指数在估算互花米草LPPC中的表现。研究表明:(1)一阶导数光谱反射率组合的植被指数用于估算互花米草的LPPC优于原始光谱反射率;(2)红边区域一阶导数光谱是估测互花米草LPPC的最佳波段;(3)对于单一色素含量的估算,叶绿素a(Chla)的最佳估算指数为FDNDVI[723,703];叶绿素b(Chlb)的最佳估算指数为FDRVI[723,525];类胡萝卜素(Cars)的最佳估算指数为FDNDVI[723,703];(4)对于使用统一参量同时估算Chla、Chlb、Cars,由FDRVI[723,703]建立的对数估算模型效果最佳。研究成果可为湿地植物生化参量反演提供参考,也可为闽江河口湿地入侵种互花米草的动态监测和生态评估管理提供有力的科学依据。     

Optical properties and the content of photosynthetic pigments in the stems and leaves of the apple-tree

Optical properties and changes in the content of photosynthetic pigments (chlorophyll and total carotenoids) were investigated in the bark and leaves of the apple-tree during a year. Optical properties of stems change with their age. Light reflectance of current year stems equalled 14.2%, while the one for 3-year-old stems decreased to 10.2%, absorption for the current year stems equalled 55.5% and increased up to 66.4% for 3-year-old ones. Light transmittance for the cork of current year stems equalled 30.2%, and decreased with the age of stems reaching 23.4% for the 3-year-old ones. The cork transmitted less than 5% of light of 400 nm, but the transmittance increased with the increase in the wavelength up to 55% at 700 nm. The reflectance of light by the leaf equalled 6.9%, absorption 89.7%, and transmittance 3.4%. In August the highest amount of chlorophyll pigments (6.2 mg·dm⁻²) and carotenoids (1.63 mg·dm⁻²) was detected in the leaves of the apple-tree, however, the ratio of chl a/b reached the highest value 4.12 in June. For the bark of apple-tree stems the content of chlorophyll pigments increased since spring and reached the maximum content of about 2.8 mg(chl)·dm⁻² for 1-3-year-old stems in the summer months, while for the current year stems in October. The ratio chl a/b was at the same level, about 2.2 during the whole year. The content of carotenoids was lower in stems than in leaves and was at the similar level during the year, however, it increased with the age of stems. Minor changes in the optical properties and the content of photosynthetic pigments occurring with the age of stems may be due to the low increment in cork thickness in the studied age groups of plants.     

Hyperspectral Remote Sensing of Canopy Biodiversity in Hawaiian Lowland Rainforests

Mapping biological diversity is a high priority for conservation research, management and policy development, but few studies have provided diversity data at high spatial resolution from remote sensing. We used airborne imaging spectroscopy to map woody vascular plant species richness in lowland tropical forest ecosystems in Hawai’i. Hyperspectral signatures spanning the 400–2,500 nm wavelength range acquired by the NASA Airborne Visible and Infrared Imaging Spectrometer (AVIRIS) were analyzed at 17 forest sites with species richness values ranging from 1 to 17 species per 0.1–0.3 ha. Spatial variation (range) in the shape of the AVIRIS spectra (derivative reflectance) in wavelength regions associated with upper-canopy pigments, water, and nitrogen content were well correlated with species richness across field sites. An analysis of leaf chlorophyll, water, and nitrogen content within and across species suggested that increasing spectral diversity was linked to increasing species richness by way of increasing biochemical diversity. A linear regression analysis showed that species richness was predicted by a combination of four biochemically-distinct wavelength observations centered at 530, 720, 1,201, and 1,523 nm (r ² = 0.85, p < 0.01). This relationship was used to map species richness at approximately 0.1 ha resolution in lowland forest reserves throughout the study region. Future remote sensing studies of biodiversity will benefit from explicitly connecting chemical and physical properties of the organisms to remotely sensed data.     

Variation in ultraviolet reflectance by carotenoid-bearing feathers of tanagers (Thraupini: Emberizinae: Passeriformes)

Avian visual sensitivity encompasses both the human visible range (400–700 nm) and also near‐ultraviolet (UV) wavelengths (320–400 nm) invisible to normal humans. I used reflectance spectrophotometry to assess variation in UV reflectance for yellow, orange and red plumage in 67 species of tanager (Passeriformes). Previous chemical studies, and my analysis of reflectance minima, suggest that carotenoids are the dominant pigments in yellow, orange and red tanager plumage. Spectra recorded over the range of wavelengths to which birds are sensitive (320–700 nm) were invariably bimodal, with both a plateau of high reflectance at longer (> 500 nm) wavelengths and a distinct secondary peak at UV (< 400 nm) wavelengths. Within this overall framework, variation in UV reflectance was expressed within well‐defined quantitative limits: (1) peak reflectance was always lower than the corresponding plateau of reflectance at longer visible wavelengths; (2) the intensity of peak reflectance declined steadily below 350 nm; (3) wavelengths of peak reflectance clustered between 350 and 370 nm. Significant correlations were detected between various measures of total reflectance in the UV and visible wavebands, but not between various measures of spectral location of UV and visible reflectance. I propose that the strong absorption band at short visible wavelengths (～ 380–550 nm) responsible for bimodal spectra of carotenoids in vitro is also responsible for bimodal reflectance by carotenoid‐based plumage colours. The construction of the UV and visible reflectance bands from different sides of this same absorbance band provides a mechanism for the observed covariation between UV and visible wavelengths. Lack of an association between the spectral locations of the UV and visible reflectance bands may result from the limited variation in spectral location of the UV band. These patterns suggest that plumage colours are subject to constraints, just as are more traditional morphological characters. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 243–257.     

Spectral reflectance indices and pigment functions during leaf ontogenesis in six subtropical landscape plants

Pigment combinations are regulated during leaf ontogenesis. To better understand pigment function, alterations in chlorophyll, carotenoid and anthocyanin concentrations were investigated during different leaf development stages in six subtropical landscape plants, namely Ixora chinensis Lam, Camellia japonica Linn, Eugenia oleina Wight, Mangifera indica L., Osmanthus fragrans Lowr and Saraca dives Pierre. High concentrations of anthocyanin were associated with reduced chlorophyll in juvenile leaves. As leaves developed, the photosynthetic pigments (chlorophyll and carotenoid) of all six species increased while anthocyanin concentration declined. Chlorophyll fluorescence imaging of Φ_PSII (effective quantum yield of PSII) and of NPQ (non-photochemical fluorescence quenching) and determination of electron transport rate-rapid light curve (RLC) showed that maximum ETR (leaf electron transport rate), Φ_PSII and the saturation point in RLC increased during leaf development but declined as they aged. Juvenile leaves displayed higher values of NPQ and Car/Chl ratios than leaves at other developmental stages. Leaf reflectance spectra (400–800 nm) were measured to provide an in vivo non-destructive assessment of pigments in leaves during ontogenesis. Four reflectance indices, related to pigment characters, were compared with data obtained quantitatively from biochemical analysis. The results showed that the ARI (anthocyanin reflectance index) was linearly correlated to anthocyanin concentration in juvenile leaves, while a positive correlation of Chl NDI (chlorophyll normalized difference vegetation index) to chlorophyll a concentration was species dependent. Photosynthetic reflectance index was not closely related to Car/Chl ratio, while a structural-independent pigment index was not greatly altered by leaf development or species. Accordingly, it is suggested that the high concentration of anthocyanin, higher NPQ and Car/Chl ratio in juvenile leaves are important functional responses to cope with high radiation when the photosynthetic apparatus is not fully developed. Another two leaf reflectance indices, ARI and Chl NDI, are valuable for in vivo pigment evaluation during leaf development.     

Surface reflectance properties of Antarctic moss and their relationship to plant species, pigment composition and photosynthetic function

In this study the variations in surface reflectance properties and pigment concentrations of Antarctic moss over species, sites, microtopography and with water content were investigated. It was found that species had significantly different surface reflectance properties, particularly in the region of the red edge (approximately 700 nm), but this did not correlate strongly with pigment concentrations. Surface reflectance of moss also varied in the visible region and in the characteristics of the red edge over different sites. Reflectance parameters, such as the photochemical reflectance index (PRI) and cold hard band were useful discriminators of site, microtopographic position and water content. The PRI was correlated both with the concentrations of active xanthophyll‐cycle pigments and the photosynthetic light use efficiency, F_v/F_m, measured using chlorophyll fluorescence. Water content of moss strongly influenced the amplitude and position of the red‐edge as well as the PRI, and may be responsible for observed differences in reflectance properties for different species and sites. All moss showed sustained high levels of photoprotective xanthophyll pigments, especially at exposed sites, indicating moss is experiencing continual high levels of photochemical stress.     

Comparative analysis of field and laboratory spectral reflectances of benthic diatoms with a modified Gaussian model approach

Benthic diatom biomass on an intertidal mudflat was estimated by field spectrometry, a non-intrusive optical method operating in the visible-infrared wavelength range. Spectral reflectance (400-900 nm) of natural assemblages was related to the amount of principal photosynthetic and accessory pigments measured by high-performance liquid chromatography (HPLC) in the top 2 mm of sediment. Relationships established in situ were compared with those obtained in the laboratory with monospecific cultures. However, to compare both types of reflectance spectra differing in their overall shape (continuum) and to isolate the pigment absorption features from other effects, we applied hyperspectral data processing, the Modified Gaussian Model (MGM), to remove the continuum and model the main absorption bands by a succession of Gaussian curves. The Gaussian band depths associated with the absorption by chlorophyll a and the pigments specific to diatoms (chlorophyll c, fucoxanthine, diadinoxanthin) displayed linear relationships with the logarithm of chlorophyll a.The most relevant spectral feature to quantify the microalgal biomass in situ was the 632 nm absorption band associated with chlorophyll c. A significant but equivocal statistical relationship was obtained at 675 nm, due to the overlapping absorption by chlorophyll breakdown products (pheophytine a and pheophorbide a) present in all the samples analyzed in the field. Fucoxanthin absorption at 550 nm can become an indicator of chlorophyll a biomass for benthic diatoms, but the effect of seasonal photophysiological adaptation should be considered. The comparative analysis of field and laboratory reflectance showed that the chlorophyll a present in the top 2 mm was not a good estimator of the photosynthetic active biomass (PAB) and that a shallower depth should be sampled in muddy sediments colonized by microphytobenthos. The approach of MGM band depth retrieval offers the possibility of quantifying PAB and has the potential to characterize (by accessory pigments) microalgae present in the photic zone of different sediment types, removing albedo variability due to grain-size scattering effects or sediment moisture content. Its accuracy relies, however, on an appropriate ground-truth sampling, excluding the contribution of redundant pigments not detected by the sensor.     

Predicting vegetation water content in wheat using normalized difference water indices derived from ground measurements

Vegetation water content (VWC) is an important variable for both agriculture and forest fire management. Remote sensing technology offers an instantaneous and non-destructive method for VWC assessment provided we can relate in situ measurements of VWC to spectral reflectance in a reliable way. In this paper, based on radiative transfer models, three new normalized difference water indices (NDWI) are proposed for VWC [fuel moisture content (FMC), and equivalent water thickness (EWT)] estimation, taking both leaf internal structure and dry matter content into account. Reflectance at 1,200, 1,450 and 1,940 nm were selected and normalized with reflectance at 860 nm to establish three water indices, NDWI₁₂₀₀, NDWI₁₄₅₀ and NDWI₁₉₄₀. Good correlations were observed between FMC (R ² = 0.65–0.80) and EWT (both at the leaf scale, R ² = 0.75–0.81 for EWT_L and at the canopy scale, R ² = 0.80–0.83 for EWT_C) at various stages of wheat crop development.     

SPECTRAL LIGHT ABSORPTION BY INTACT BLADES OF PORPHYRA ABBOTTAE (RHODOPHYTA): EFFECTS OF ENVIRONMENTAL FACTORS IN CULTURE1

Whole thallus absorptance spectra were recorded for Porphyra abbottae Krishnamurthy gametophytes grown in batch culture at combinations of temperature (8, 10, 12° C), irradiance (17.5, 70, 140 μmol photons·m^?2·s^?1), nutrients (f/4, f/2, f media) and water motion (0, 50, 100, 150 rpm). Light, nutrients, water motion and the interaction of nutrients with water motion all significance affected broadband (400-700 nm) absorptance and absorptance by phycoerythrin (566 nm), phycocyanin (624 nm) and chlorophyll a (680 nm). Absorptances increased in low light, low water motion and high nutrient levels. Shifts in phycoerythrin: chlorophyll a absorptance ratios closely paralleled changes of absorptance by the major pigments, whereas the phycoerythrin: phycocyanin ratio decreased only with increasing nutrient supply Absorptance ratios were significantly correlated with growth rate. Absorptance increased asymptotically with blade thickness or pigment content. Based on previously determined growth rates, nutrient saturated P. abbottae can synthesize photosynthetic pigments in excess of immediate needs. Allocation is given preferentially to the phycobiliproteins, with highest preference for phycocyanin.     

Spectral reflectance of coral

Spectral reflectance (R) of corals is a fundamental parameter to coral reef remote sensing. We explore general trends as well as geographic and taxonomic variabilities of coral R using a data set consisting of 5,199 Rs measured in situ at depths of up to 15 m for 195 coral colonies at 11 sites worldwide. Coral R ranges in magnitude from ~0.5% at 400 nm to near 100% at 700 nm; mean coral R rises from ~2.5% at 400–500 nm to ~8% between 550 and 650 nm. All corals measured in this study exhibit one of two basic shapes of R, which we label the brown and blue modes. We postulate that brown-mode R is determined by pigment absorption solely by zooxanthellae, while blue-mode R arises through expression of a non-fluorescing coral-host pigment. Taxonomic and geographic variabilities are approximately equal to global variability, both in magnitude and shape, indicating that coral R is independent of taxonomic or geographic differences. We reason that this is to be expected, since R is determined by pigments that are conservative across geographic and taxonomic boundaries.     

Leaf characteristics and optical properties of different woody species

 Light partition has been examined and evaluated on five woody species (Olea europaea, Ficus carica, Pittosporum tobira, Hedera helix maculata, Persica vulgaris) in relation to their leaf morpho-histological characteristics, water and chlorophyll contents. Leaf parameters and optical properties (reflectance, transmittance, absorbance) in PAR, FR and NIR wavebands (400–1100 nm) were preliminarily submitted to a canonical correlation analysis where lamina thickness and water content showed a leading role in determining all the optical properties, while chlorophyll, influential in the PAR region, was remarkably effective only in an extreme pigment situation when green and albino patches of ivy leaves were compared. Transmittance appeared inversely related to lamina thickness in accordance with the Lambert Beer law. Significant correlations were found also between mesophyll water content and both transmittance (positive) and reflectance (negative). Olive leaves showed peculiar optical patterns because of the dense and continuous trichome layer on their abaxial surface. Received: 3 January 1997 / Accepted: 5 May 1997     

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。     

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。     

Effects of lifelong [CO2] enrichment on carboxylation and light utilization of Quercus pubescens Willd. examined with gas exchange,biochemistry and optical techniques

Lifelong exposure to elevated concentrations of atmospheric CO₂ may enhance carbon assimilation of trees with unlimited rooting volume and consequently may reduce requirements for photoprotective pigments. In early summer the effects of elevated [CO₂] on carboxylation and light utilization of mature Quercus pubescens trees growing under chronic [CO₂] enrichment at two CO₂ springs and control sites in Italy were examined. Net photosynthesis was enhanced by 36 to 77%. There was no evidence of photosynthetic downregulation early in the growing season when sink demand presumably was greatest. Specifically, maximum assimilation at saturating [CO₂], electron transport capacity, and Rubisco content, activity and carboxylation capacity were not significantly different in trees growing at the CO₂ springs and their respective control sites. Foliar biochemical content, leaf reflectance index of chlorophyll pigments (NDVI), and photochemical efficiency of PSII (ΔF/F_m′) also were not significantly affected by [CO₂] enrichment except that starch content and ΔF/F_m′ tended to be higher at one spring (42 and 15%, respectively). Contrary to expectation, prolonged elevation of [CO₂] did not reduce xanthophyll cycle pigment pools or alter mid‐day values of leaf reflectance index of xanthophyll cycle pigments (PRI), despite the enhancement of carbon assimilation. However, both these pigments and PRI were well correlated with electron transport capacity.     

Salt impact on photosynthesis and leaf ultrastructure of <Emphasis Type="Italic">Aeluropus littoralis</Emphasis>

The effects of salinity (400 mM NaCl) on growth, biomass partitioning, photosynthesis, and leaf ultrastructure were studied in hydroponically grown plants of Aeluropus littoralis (Willd) Parl. NaCl produced a significant inhibition of the main growth parameters and a reduction in leaf gas exchange (e.g. decreased rates of photosynthesis and stomatal conductance). However, NaCl salinity affected neither the composition of photosynthesis pigments nor leaf water content. The reduction in leaf gas exchange seemed to correlate with a decrease in mesophyll thickness as well as a severe disorganisation of chloroplast structure, with misshapen chloroplasts and dilated thylakoid membranes. Conspicuously, mesophyll chloroplasts were more sensitive to salt treatment than those of bundle sheath cells. The effects of NaCl toxicity on leaf structure and ultrastructure and the associated physiological implications are discussed in relation to the degree of salt resistance of A. littoralis.     

Reflectance Variation within the In-Chlorophyll Centre Waveband for Robust Retrieval of Leaf Chlorophyll Content

The in-chlorophyll centre waveband (ICCW) (640–680 nm) is the specific chlorophyll (Chl) absorption band, but the reflectance in this band has not been used as an optimal index for non-destructive determination of plant Chl content in recent decades. This study develops a new spectral index based solely on the ICCW for robust retrieval of leaf Chl content for the first time. A glasshouse experiment for solution-culture of one chlorophyll-deficient rice mutant and six wild types of rice genotypes was conducted, and the leaf reflectance (400–900 nm) was measured with a high spectral resolution (1 nm) spectrophotometer and the contents of chlorophyll a (Chla), chlorophyll b (Chlb) and chlorophyll a+b (Chlt) of the rice leaves were determined. It was found that the reflectance curves from 640 nm to 674 nm and from 675 nm to 680 nm of the low-chlorophyll mutant leaf were drastically steeper than that of the wild types in the ICCW. The new index based on the reflectance variation within ICCW, the difference of the first derivative sum within the ICCW (DFDS_ICCW), was highly sensitive (r = −0.77, n = 93, P<0.01) to Chlt while the mean reflectance (R_ICCW) in the ICCW became insensitive (r = −0.12, n = 93, P>0.05) to Chlt when the leaf Chlt was higher than 200 mg/m². The best equations of R-ICCW and DFDS_ICCW yielded an RMSE of 78.7, 32.9 and 107.3 mg/m², and an RMSE of 37.4, 16.0 and 45.3 mg/m⁻², respectively, for predicting Chla, Chlb and Chlt. The new index could rank in the top 10 for prediction of Chla and Chlt as compared with the 55 existing indices. Additionally, most of the 55 existing Chl-related VIs performed robustly or strongly in simultaneous prediction of leaf Chla, Chlb and Chlt.     

Altitudinal differences in UV absorbance,UV reflectance and related morphological traits of Quercus ilex and Rhododendron ferrugineum in the Mediterranean region

We studied the variations in different physiological parameters associated with UV-B radiation defense: UV-B radiation absorbance, UV-visible spectral reflectance, carotenoids concentration, leaf thickness, SLW (specific leaf weigth) and trichome density in Quercus ilex growing at 200 and 1200 m and Rhododendron ferrugineum growing at 2200 m. We examined the role of these parameters as protection mechanisms in an altitudinal gradient of increasing UV radiation in northern Catalonia and in sun and shade leaves. The concentration of UV-B radiation absorbing pigments was 15% higher in sun leaves of Q. ilex at 1200 m than in those from 200 m altitude. Sun leaves of R. ferrugineum presented concentrations three times higher than those of Q. ilex. Reflectance ranged between 5% (in the region 300–400 nm) and 12% (in the region 280–300 nm). The variation of reflectance with altitude followed an inverse trend respect to absorbance in the 280–300 nm region, with higher values the lower the altitude, but in the 300–400 nm region, reflectance of the lower site was the lowest. In both species and altitudes sun leaves presented higher concentrations of UV-B radiation absorbing pigments and UV reflectance than shade leaves. Quercus ilex trees of the higher location presented higher NDPI (Normalized Difference Pigment Reflectance Index) values, indicating higher carotenoids/chlorophyll a ratio. Actual measurements of carotenoid/chlorophyll a ratio confirmed this pattern. The photochemical reflectance index (PRI) presented higher values the higher the location indicating lower photosynthetic radiation-use efficiency. Specific leaf weight (SLW) and leaf thickness were larger in Q. ilex trees of higher location than in those of lower location. In both sites, sun leaves also presented larger SLW values than shade leaves. Adaxial leaf hair density in sun leaves was significantlly higher in the lower location. UV absorption and linked morphological traits (SLW, leaf thickness measured in Q. ilex) presented the larger differences among studied plants at different altitudes and seem to be the dominant UV protecting mechanisms.     

Contribution of intercellular reflectance to photosynthesis in shade leaves

The potential contribution of intercellular light reflectance to photosynthesis was investigated by infiltrating shade leaves with mineral oil. Infiltration of leaves of Hydrophyllum canadense and Asarum canadense with mineral oil decreased adaxial leaf reflectance but increased transmittance. As a result of the large increase in transmittance, infiltration caused a decrease in absorptance of 25% and 30% at 550 and 750 nm, respectively. Thus, intercellular reflectance increased absorptance in these species by this amount. In a comparison of sun and shade leaves of Acer saccharum and Parthenocissus quinquefolia, oil infiltration decreased absorptance more in shade than in sun leaves. This difference suggests that the higher proportion of spongy mesophyll in shade leaves may increase internal light scattering and thus absorptance. The importance of the spongy mesophyll in increasing internal reflectance was also evident in comparisons of the optics of Populus leaves and in the fluorescence yield of oil-infiltrated leaves of several sun and shade species. Oil infiltration decreased the quantum yield of fluorescence (F_o) by 39–52% for shade leaves but only 21–25% for sun leaves. We conclude that the greater proportion of spongy parenchyma in shade leaves increased intercellular light scattering and thus absorptance. Direct measurements with fibre-optic light probes of the distribution of light inside leaves of Hydrophyllum canadense confirmed that oil infiltration decreased the amount of back-scattered light and that most of the light scattering for this species occurred from the middle of the palisade layer to the middle of the spongy mesophyll. We were not, however, able to assess the potential contribution of reflectance from the internal abaxial epidermis to total internal light scattering in these experiments. Using a mathematical model to compare the response of net photosynthesis (O₂, flux) to incident irradiance for control leaves of H. canadense and theoretical leaves with no intercellular reflectance, we calculated that intercellular reflectance caused a 1.97-fold increase in photosynthesis at 20 μmol m^?2s^?1 (incident photon flux density). This enhancement of absorption and photosynthesis by inter-cellular reflectance, without additional production and maintenance of photosynthetic pigments, may maintain shade leaves above the photosynthetic light compensation point between sunflecks and maintain the light induction state during protracted periods of low diffuse light.     

Spectroscopic and Biochemical Analyses of UV Effects on Phycobiliproteins of Anabaena sp. and Nostoc carmium

The effects of UV (280–400 nm) irradiation on phycobiliprotein composition have been studied in two N₂-fixing cyanobacteria, Anabaena sp. and Nostoc carmium, isolated from rice paddy fields in India. Phycobiliproteins were isolated and separated by sucrose density gradient centrifugation. After UV exposure the top fraction mainly contained carotenoids (absorption maximum at 485 nm), which first showed an increase in intensity and absorption and then a gradual decrease with increasing UV exposure in Anabaena sp., whereas, in Nostoc carmium this fraction showed a steady increase over the whole exposure time. The bottom fraction of both organisms mainly contained phycocyanin (absorption peak at 620 nm) which showed a steady decline in intensity, as well as absorption. Fluorescence excitation at 620 nm resulted in an emission at 650 nm which underwent a shift towards shorter wave-lengths with increasing UV-exposure time, indicating a disassembly of the phycobilisomal complex and of impaired energy transfer from accessory pigments to the reaction centers. SDS PAGE analysis of the fractions revealed a loss of high molecular mass linker proteins and low molecular mass (αβ monomers indicating that the phycobiliproteins, which function as accessory pigments for the operation of photosystem II, disassemble during UV irradiation.