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.     

Epidermis effects on spectral properties of leaves of four herbaceous species

The spectral properties of the leaves of the herbaceous species Brassica oleracea L. var. botrytis L., Cerastum tomentosum L., Petunia hybrida Vilm., and Talinum paniculatum (Jacq.) Gaertn. were examined to see what effect the epidermis had on leaf absorptance, reflectance and transmittance. Removal of the epidermis from the side of the leaf surface being illuminated resulted in increases in leaf absorptance and transmittance, and a decrease in reflectance in the 400–800 nm waveband. Removal of the epidermis from the opposite side of an illuminated leaf (effect was similar in both abaxial and adaxial surfaces) resulted in small decreases in both absorptance and reflectance, and corresponding increases in transmittance. Removal of both the upper and lower epidermis resulted in a marked increase in transmittance, while both leaf reflectance and absorptance were decreased. The results suggest that the presence of the epidermis significantly increases leaf absorptance in the photosynthetic wavebands.     

The content of photosynthetic pigments and the light conditions in the fruits and leaves of sweet pepper

Investigations were carried out on the fruits of sweet pepper at its two development stages: on green fruits, on mature red and yellow fruits and on leaves. The content of the photosynthetic pigments and the optical properties were examined. In the green fruits when compared with leaves the content of the photosynthetic pigments is smaller by 40 to 50 % and the value of the ratio: chlorophyll a/b is lower. Chlorophyll is absent in mature fruits, while the content of carotenoids is a few times higher. The optical properties of green fruits and of the leaves in the PAR (photosynthetically active radiation) range are the same. In the range 700–1100 nm the green fruits show smaller reflectance and transmittance and a few times greater absorptance of irradiation which contributes to the warming up of the seed bag, while small absorptance of leaves in this range protects them against overheating. In mature fruits, in the PAR range, the reflectance and transmittance are higher, while the absorptance of irradiation in comparison with that of green fruits is smaller. In the range 700 – 1100 nm the changes are rather small and refer to the increase of reflectance and reduction of absorptance, while the transmittance of irradiation remains unchanged.     

Effect of drought stress on leaf optical properties in drought-resistant and drought-sensitive maize and triticale genotypes

The effect of a short (7 d), prolonged (14 d) soil drought (D) and (7 d) recovery (DR) on the leaf optical properties — reflectance (R), transmittance (T) and absorptance (A) in photosynthetically active radiation (PAR) and near infrared radiation (NIR) range of irradiation (750–1100 nm) was studied for maize and triticale genotypes differing in drought tolerance. The drought stress caused the changes in leaf optical properties parameters in comparison with non-drought plants. The observed harmful influence of drought was more visible for maize than triticale.     

LEAF OPTICAL PROPERTIES OF RAINFOREST SUN AND EXTREME SHADE PLANTS

The optical properties of the leaves of twelve tropical sun species and thirteen tropical extreme shade species were examined with an integrating sphere attached to a spectroradiometer. Measurements of diffuse reflectance and transmittance allowed calculations of absorptance, 350–1,100 nm. Although some shade species absorbed higher percentages of quantum flux densities for photosynthesis (400–700 nm, PPFD) than the mean for the sun species, the sun and shade species as groups were not significantly different from each other: 90.2, S.D. 3.6% for shade species and 88.6, S.D. 2.4% for the sun species. The groups of species did not differ in total absorptance of energy 350–1,100 nm. Furthermore, the sun and shade species were identical in their shift of absorptance at wavelengths between 650 and 750 nm. The anthocyanic coloration of the leaf undersurfaces of two species polymorphic for this characteristic (Trionela hirsuta and Ischnosciphon pruinosus) is correlated with increased absorptance at the upper end of the action spectrum of photosynthesis. Although sun and shade species have similar optical properties, the energy investment (as documented by dry wt per unit area of leaf surface) is much less for the shade species.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

固定对组织光学性质的影响

利用带有积分球的SHIMADZU UV—240分光光度计,测量了组织在固定前和后的反射率和透射率,分析了福尔马林固定对组织光学性质的影响。     

LEAF OPTICAL PROPERTIES ALONG A VERTICAL GRADIENT IN A TROPICAL RAIN FOREST CANOPY IN COSTA RICA

Leaf optical properties (400–1,100 nm) were compared for four species of rain forest trees with crowns in understory, mid-canopy, and canopy positions to test whether optical properties change with light environment. The species tested represent a spectrum of regeneration patterns ranging from shade tolerant to light demanding. Overall, leaf optical properties of the four species were similar. Differences in absorptance were small, but statistically significant among the species and positions along the canopy gradient. Species absorptance differences corresponded somewhat to shade tolerance; two of the shade species showed higher absorptance in lower light environments, while the sun species showed the reverse pattern. Specific leaf mass (leaf weight per unit area) and chlorophyll content per unit leaf weight also changed along the canopy gradient. Specific leaf mass was positively correlated and chlorophyll per unit leaf weight was negatively correlated with increasing light environment. Consequently, the efficiency of absorption, as represented by the absorptance per unit leaf weight, increased as light level decreased, largely due to changes in specific leaf mass. In contrast, efficiency of absorption per unit leaf chlorophyll was relatively constant with light environment for the two species measured for chlorophyll.     

The combined effects of CO2 concentration and enhanced UV-B radiation on faba bean. 3. Leaf optical properties,pigments, stomatal index and epidermal cell density

Seedlings of Vicia faba L. (cv. Minica) were grown in a factorial experiment in a greenhouse. The purpose of the study was to determine whether CO₂ enrichment and supplemental UV-B radiation affect leaf optical properties and whether the combined effects differ from single factor effects. Seedlings were grown at either 380 mol mol^-1 or 750 mol mol^-1 CO₂ and at four levels of UV-B radiation. After 20 and 40 days of treatment, absorptance, transmittance and reflectance of photosynthetically active radiation (PAR) were measured on the youngest fully developed leaf. On the same leaf, the specific leaf area on a fresh weight basis (SLA_fw), chlorophyll content, UV-B absorbance, transmittance of UV light and stomatal index were measured. UV-B radiation significantly increased PAR absorptance and decreased PAR transmittance. The increased PAR absorptance can be explained by an increased chlorophyll content in response to UV-B radiation. Leaf transmittance of UV radiation decreased with increasing UV-B levels mainly caused by increased absorbance of UV absorbing compounds. UV-B radiation decreased both the stomatal density and epidermal cell density of the abaxial leaf surface, leaving the stomatal index unchanged. Effects of CO₂ enrichment were less pronounced than those of UV-B radiation. The most important CO₂ effect was an increase in stomatal density and epidermal cell density of the adaxial leaf surface. The stomatal index was not affected. No interaction between CO₂ and UV-B radiation was found. The results are discussed in relation to the internal light environment of the leaf.     

Variability in leaf optical properties among 26 species from a broad range of habitats

Leaves from 26 species with growth forms from annual herbs to trees were collected from open, intermediate, and shaded understory habitats in Mississippi and Kansas, USA. Leaf optical properties including reflectance, transmittance, and absorptance in visible and near infrared (NIR) wavelengths were measured along with leaf thickness and specific leaf mass (SLM). These leaf properties and internal light scattering have been reported to vary with light availability in studies that have focused on a limited number of species. Our objective was to determine whether these patterns in leaf optics and light availability were consistent when a greater number of species were evaluated. Leaf thickness and SLM varied by tenfold among species sampled, but within-habitat variance was high. Although there was a strong trend toward thicker leaves in open habitats, only SLM was significantly greater in open vs. understory habitats. In contrast, leaf optical properties were strikingly similar among habitats. Reflectance and reflectance/transmittance in the NIR were used to estimate internal light scattering and there were strong relationships (r1 > 0.65) between these optical properties and leaf thickness. We concluded that leaf thickness, which did not vary consistently among habitats, was the best predictor of NIR reflectance and internal light scattering. However, because carbon allocation to leaves was lower in understory species (low SLM) yet gross optical properties were similar among all habitats, the energy investment by shade leaves required to achieve optical equivalence with sun leaves was lower. Differences in leaf longevity and growth form within a habitat may help explain the lack of consistent patterns in leaf optics as the number of species sampled increases.     

Elevated CO2 interacts with herbivory to alter chlorophyll fluorescence and leaf temperature in Betula papyrifera and Populus tremuloides

Herbivory can influence ecosystem productivity, but recent evidence suggests that damage by herbivores modulates potential productivity specific to damage type. Because productivity is linked to photosynthesis at the leaf level, which in turn is influenced by atmospheric CO(2) concentrations, we investigated how different herbivore damage types alter component processes of photosynthesis under ambient and elevated atmospheric CO(2). We examined spatial patterns in chlorophyll fluorescence and the temperature of leaves damaged by leaf-chewing, gall-forming, and leaf-folding insects in aspen trees as well as by leaf-chewing insects in birch trees under ambient and elevated CO(2) at the aspen free-air CO(2) enrichment (FACE) site in Wisconsin. Both defoliation and gall damage suppressed the operating efficiency of photosystem II (ΦPSII) in remaining leaf tissue, and the distance that damage propagated into visibly undamaged tissue was marginally attenuated under elevated CO(2). Elevated CO(2) increased leaf temperatures, which reduced the cooling effect of gall formation and freshly chewed leaf tissue. These results provide mechanistic insight into how different damage types influence the remaining, visibly undamaged leaf tissue, and suggest that elevated CO(2) may reduce the effects of herbivory on the primary photochemistry controlling photosynthesis.     

Using leaf optical properties to detect ozone effects on foliar biochemistry

Efficient methods for accurate and meaningful high-throughput plant phenotyping are limiting the development and breeding of stress-tolerant crops. A number of emerging techniques, specifically remote sensing methods, have been identified as promising tools for plant phenotyping. These remote sensing methods can be used to accurately and rapidly relate variations in leaf optical properties with important plant characteristics, such as chemistry, morphology, and photosynthetic properties at the leaf and canopy scales. In this study, we explored the potential to utilize optical (λ = 500–2,400 nm) near-surface remote sensing reflectance spectroscopy to evaluate the effects of ozone pollution on photosynthetic capacity of soybean (Glycine max Merr.). The research was conducted at the Soybean Free Air Concentration Enrichment (SoyFACE) facility where we subjected plants to ambient (44 nL L^?1) and elevated ozone (79–82 nL L^?1 target) concentrations throughout the growing season. Exposure to elevated ozone resulted in a significant loss of productivity, with the ozone-treated plants displaying a ~30 % average decrease in seed yield. From leaf reflectance data, it was also clear that elevated ozone decreased leaf nitrogen and chlorophyll content as well as the photochemical reflectance index (PRI), an optical indicator of the epoxidation state of xanthophyll cycle pigments and thus physiological status. We assessed the potential to use leaf reflectance properties and partial least-squares regression (PLSR) modeling as an alternative, rapid approach to standard gas exchange for the estimation of the maximum rates of RuBP carboxylation (V _c,max), an important parameter describing plant photosynthetic capacity. While we did not find a significant impact of ozone fumigation on V _c,max, standardized to a reference temperature of 25 °C, the PLSR approach provided accurate and precise estimates of V _c,max across ambient plots and ozone treatments (r ² = 0.88 and RMSE = 13.4 μmol m^?2 s^?1) based only on the variation in leaf optical properties and despite significant variability in leaf nutritional status. The results of this study illustrate the potential for combining the phenotyping methods used here with high-throughput genotyping methods as a promising approach for elucidating the basis for ozone tolerance in sensitive crops.     

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     

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.     

Changes in the optical properties of cotyledons of Cucurbits pepo during the first seven days of their development

Abstract. We recently discussed a method for measuring optical properties of light scattering and absorbing plant tissue ( Seyfried, Fukshansky & Schafer, 1983 ). This method has been used to measure the changes in optical properties of cotyledons between 360 and 1000 run during the first 7d of development. The seedlings were either etiolated or grown under continuous white light, the latter either herbicide-treated (SAN 9789 = Norflurazon) or untreated. Some of the observed changes in seedlings grown under white light are due to chlorophyll accumulation. This accumulation leads to an increase in absorption coefficients at all wavelengths except in the 750 to 850 nm region. Reflectance, transmittance, and the scattering coefficient decreased markedly. Other changes seem to be independent of light conditions since they occur in much the same way under all treatments. These are a generally decreasing reflectance and scattering coefficient and an even stronger decrease of reflectance from the upper face of the cotyledon as compared to the reflectance from the lower face, in particular in the blue region of the spectrum. The observed changes are discussed in terms of light gradients and the resulting problems for in vivo spectroscopy.     

Leaf optical responses to light and soil nutrient availability in temperate deciduous trees

Leaf optical parameters influence light availability at the cellular, leaf, and canopy scale of integration. While recent studies have focused on leaf optical responses to acute plant stress, the effects of changes in plant resources on leaf optics remain poorly characterized. We examined leaf optical and anatomical responses of five temperate deciduous tree species to moderate changes in nutrient and light availability. Spectral reflectance in the visible waveband generally increased at high light, but decreased with increased nutrient availability. Patterns of both spectral reflectance and absorptance were primarily determined by chlorophyll concentration although carotenoid concentration was also influential. While most anatomical features did not explain residual variation in reflectance, cuticle thickness was significantly related to reflectance at complementary angles compared to the angle of incidence. Absorptance did not change with light environment; however, absorption efficiency per unit biomass increased by approximately 40% under low light, due to reduced leaf mass per area. We conclude that changes in resource availability differentially influence leaf optical properties and that such changes are driven primarily by changes in pigment concentrations. The magnitude of leaf optical responses to moderate changes in resource availability was comparable to those of acute stress responses and varied among species.     

Does elevated CO2 ameliorate the impact of O3 on chlorophyll content and photosynthesis in potato (Solanum tuberosum)?

This study examined the impact of season-long exposure to elevated carbon dioxide (CO2) and ozone (O3), individually and in combination, on leaf chlorophyll content and gas exchange characteristics in potato (Solanum tuberosum L. cv. Bintje). Plants grown in open-top chambers were exposed to three CO2 (ambient, 550 and 680 μmol mol-1) and two O3 treatments (ambient and elevated; 25 and 65 nmol mol-1, 8 h day-1 means, respectively) between crop emergence and maturity; plants were also grown in unchambered field plots. Non-destructive measurements of chlorophyll content and visible foliar injury were made for all treatments at 2-week intervals between 43 and 95 days after emergence. Gas exchange measurements were made for all except the intermediate 550 μmol mol-1 CO2 treatment. Season-long exposure to elevated O3 under ambient CO2 reduced chlorophyll content and induced extensive visible foliar damage, but had little effect on net assimilation rate or stomatal conductance. Elevated CO2 had no significant effect on chlorophyll content, but greatly reduced the damaging impact of O3 on chlorophyll content and visible foliar damage. Light-saturated assimilation rates for leaves grown under elevated CO2 were consistently lower when measured under either elevated or ambient CO2 than in equivalent leaves grown under ambient CO2. Analysis of CO2 response curves revealed that CO2-saturated assimilation rate, maximum rates of carboxylation and electron transport and respiration decreased with time. CO2-saturated assimilation rate was reduced by elevated O3 during the early stages of the season, while respiration was significantly greater under elevated CO2 as the crop approached maturity. The physiological origins of these responses and their implications for the performance of potato in a changing climate are discussed.     

Adaxial/abaxial specification in the regulation of photosynthesis and stomatal opening with respect to light orientation and growth with CO2 enrichment in the C4 species Paspalum dilatatum.

Whole-plant morphology, leaf structure and composition were studied together with the effects of light orientation on the dorso-ventral regulation of photosynthesis and stomatal conductance in Paspalum dilatatum cv. Raki plants grown for 6 wk at either 350 or 700 microl l(-1) CO(2). Plant biomass was doubled as a result of growth at high CO(2) and the shoot:root ratio was decreased. Stomatal density was increased in the leaves of the high CO(2)-grown plants, which had greater numbers of smaller stomata and more epidermal cells on the abaxial surface. An asymmetric surface-specific regulation of photosynthesis and stomatal conductance was observed with respect to light orientation. This was not caused by dorso-ventral variations in leaf structure, the distribution of phosphoenolpyruvate carboxylase (PEPC) and ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) proteins or light absorptance, transmittance or reflectance. Adaxial/abaxial specification in the regulation of photosynthesis results from differential sensitivity of stomatal opening to light orientation and fixed gradients of enzyme activation across the leaf.     

Changes in leaf optical properties associated with light-dependent chloroplast movements

We surveyed 24 plant species to examine how leaf anatomy influenced chloroplast movement and how the optical properties of leaves change with chloroplast position. All species examined exhibited light-dependent chloroplast movements but the associated changes in leaf absorptance varied considerably in magnitude. Chloroplast movement-dependent changes in leaf absorptance were greatest in shade species, in which absorptance changes of >10% were observed between high- and low-light treatments. Using the Kubelka-Munk theory, we found that changes in the absorption (k) and chlorophyll a absorption efficiency (k*) associated with chloroplast movement correlated with cell diameter, such that the narrower, more columnar cells found in sun leaves restricted the ability of chloroplasts to move. The broader, more spherical cells of shade leaves allowed greater chloroplast rearrangements and in low-light conditions allowed efficient light capture. Across the species tested, light-dependent chloroplast movements modulated leaf optical properties and light absorption efficiency by manipulating the package (sieve or flattening) effect but not the detour (path lengthening) effect.     

Plant ecophysiological processes in spectral profiles: perspective from a deciduous broadleaf forest

The need for progress in satellite remote sensing of terrestrial ecosystems is intensifying under climate change. Further progress in Earth observations of photosynthetic activity and primary production from local to global scales is fundamental to the analysis of the current status and changes in the photosynthetic productivity of terrestrial ecosystems. In this paper, we review plant ecophysiological processes affecting optical properties of the forest canopy which can be measured with optical remote sensing by Earth-observation satellites. Spectral reflectance measured by optical remote sensing is utilized to estimate the temporal and spatial variations in the canopy structure and primary productivity. Optical information reflects the physical characteristics of the targeted vegetation; to use this information efficiently, mechanistic understanding of the basic consequences of plant ecophysiological and optical properties is essential over broad scales, from single leaf to canopy and landscape. In theory, canopy spectral reflectance is regulated by leaf optical properties (reflectance and transmittance spectra) and canopy structure (geometrical distributions of leaf area and angle). In a deciduous broadleaf forest, our measurements and modeling analysis of leaf-level characteristics showed that seasonal changes in chlorophyll content and mesophyll structure of deciduous tree species lead to a seasonal change in leaf optical properties. The canopy reflectance spectrum of the deciduous forest also changes with season. In particular, canopy reflectance in the green region showed a unique pattern in the early growing season: green reflectance increased rapidly after leaf emergence and decreased rapidly after canopy closure. Our model simulation showed that the seasonal change in the leaf optical properties and leaf area index caused this pattern. Based on this understanding we discuss how we can gain ecophysiological information from satellite images at the landscape level. Finally, we discuss the challenges and opportunities of ecophysiological remote sensing by satellites.