A widely used algorithm, Sen2Cor, for the atmospheric correction of Sentinel-2 MSI images assumes a constant ratio of red (0.6 μm) and near-infrared (2.2 μm) spectral reflectance, taking ρ2.2/ρ0.6 = 2. In fact, however, this ratio exhibits substantial dependence on vegetation phenology. The discrepancy produces errors in predicted surface reflectance. We analyse the reflectance of 92,230 forest stands, using full time series of Landsat-8/9 OLI and Sentinel-2 MSI images taken over Estonia. We find the ρ2.2/ρ0.6 value for evergreen needleleaf forests to be rather stable over time, with a mean value around 2.25. In broadleaf deciduous forests, on the other hand, we find ρ2.2/ρ0.6 to increase rapidly in the spring, to remain on average close to 3 or greater, and then to decrease in the autumn. We offer parametric and tabulated models of ρ2.2/ρ0.6 for eventual integration with Sen2Cor.
This chapter gives a theoretical overview of various contact, proximal and remote monitoring solutions available for precision agriculture. Visual inspection of crop damage, which can be detected using these sensors, are introduced at first. Precision agriculture methodologies and sensors are reviewed with particular emphasis on variable rate fertilization. Different sensor platforms reviewed in the chapter ranged from drone images to tractor-mounted and hand-held devices, including the overview of autonomous platforms and robots in precision agriculture. After the theoretical overview a couple of use-cases are described to illustrate the most common practices of using proximal sensing sensors for precision agriculture. The use-case from Estonia demonstrates hand-held proximal sensor usage for variable rate fertilization. The use-cases from Lithuania illustrate field-scale monitoring and mapping of soil characteristics.
We examined the seasonal changes in biophysical, anatomical, and optical traits of young leaves, formed throughout the vegetative season due to sylleptic growth, and mature leaves formed by proleptic growth in spring. Leaf developmental categories contribute to the top-of-canopy reflectance and should be considered when taking ground truth for remote sensing studies (RS). Deciduous tree species, Betula pendula, Populus tremula, and Alnus incana, were sampled from May to October 2018 in an Estonian hemiboreal forest. Chlorophyll and carotenoid content were detected biochemically; leaf anatomical traits (leaf, palisade, and spongy mesophyll thickness) were measured on leaf cross-sections; leaf reflectance was measured by a spectroradiometer with an integrating sphere (350–2500 nm). Biophysical and anatomical leaf traits were related to 64 vegetation indices (VIs). Linear models based on VIs for all tested leaf traits were more robust if both juvenile and mature leaves were included. This study provides information on which VIs are interchangeable or independent. Pigment and leaf thickness sensitive indices formed PC1; water and structural trait related VIs formed an independent group associated with PC3. Type of growth and leaf age could affect the validation of biophysical and anatomical leaf trait retrieval from the optical signal. It is, therefore, necessary to sample both leaf developmental categories—young and mature—in RS, especially if sampling is only once within the vegetation season.
This study present the results of airborne top-of-canopy measurements of reflectance spectra in the spectral domain of 350–1050 nm over the hemiboreal mixed forest. We investigated spectral transformations that were originally designed for utilization at very different spectral resolutions. We found that the estimates of red edge inflection point by two methods—the linear four-point interpolation approach (S2REP) and searching the maximum of the first derivative spectrum ( D m a x ) according to the mathematical definition of red edge inflection point—were well related to each other but S2REP produced a continuously shifting location of red edge inflection point while D m a x resulted in a discrete variable with peak jumps between fixed locations around 717 nm and 727 nm for forest canopy (the third maximum at 700 nm appeared only in clearcut areas). We found that, with medium high spectral resolution (bandwidth 10 nm, spectral step 3.3 nm), the in-filling of the O 2 -A Fraunhofer line ( F a r e a ) was very strongly related to single band reflectance factor in NIR spectral region ( ρ = 0.91, p < 0.001) and not related to Photochemical Reflectance Index (PRI). Stemwood volume, basal area and tree height of dominant layer were negatively correlated with reflectance factors at both visible and NIR spectral region due to the increase in roughness of canopy surface and the amount of shade. Forest age was best related to single band reflectance at NIR region ( ρ = −0.48, p < 0.001) and the best predictor for allometric LAI was the single band reflectance at red spectral region ( ρ = −0.52, p < 0.001) outperforming all studied vegetation indices. It suggests that Sentinel-2 MSI bands with higher spatial resolution (10 m pixel size) could be more beneficial than increased spectral resolution for monitoring forest LAI and age. The new index R 751 /R 736 originally developed for leaf chlorophyll content estimation, also performed well at the canopy level and was mainly influenced by the location of red edge inflection point ( ρ = 0.99, p < 0.001) providing similar info in a simpler mathematical form and using a narrow spectral region very close to the O 2 -A Fraunhofer line.
Vegetation indices are calculated from reflectance data of discrete spectral bands. The reflectance signal in the visible spectral range is dominated by the optical properties of photosynthetic pigments in plant leaves. Numerous spectral indices have been proposed for the estimation of leaf pigment contents, but the efficacy of different indices for prediction of pigment content and composition for species-rich communities is unknown. Assessing the ability of different vegetation indices to predict leaf pigment content we identify the most suitable spectral indices from an experimental dataset consisting of field-grown high light exposed leaves of 33 angiosperm species collected in two sites in Mallorca (Spain) with contrasting leaf anatomy and pigment composition. Leaf-level reflectance spectra were recorded over the wavelength range of 400 – 900 nm and contents of chlorophyll a, chlorophyll b, total carotenoids, and anthocyanins were measured in 33 species from different plant functional types, covering a wide range of leaf structures and pigment content, five-fold to more than 10-fold for different traits. The best spectral region for estimation of leaf total chlorophyll content with least interference from carotenoids and anthocyanins was the beginning of near-infrared plateau well beyond 700 nm. Leaves of parallel-veined monocots and pinnate-veined dicots had different relationships between vegetation indices and pigments. We suggest that the nature and role of “far-red” chlorophylls which absorb light at longer wavelengths than 700 nm constitute a promising target for future remote sensing studies.
Plants can respond to climate change by either migrating, adapting to the new conditions or going extinct. Relict plant species of limited distribution can be especially vulnerable as they are usually composed of small and isolated populations, which may reduce their ability to cope with rapidly changing environmental conditions. The aim of this study was to assess the vulnerability of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub of limited distribution, to a future drier climate. We evaluated population differentiation in functional traits related to drought tolerance across seven representative populations of the species' range. We measured morphological and physiological traits in both the field and the greenhouse under three water availability levels. Large phenotypic differences among populations were found under field conditions. All populations responded plastically to simulated drought, but they differed in mean trait values as well as in the slope of the phenotypic response. Particularly, dry-edge populations exhibited multiple functional traits that favored drought tolerance, such as more sclerophyllous leaves, strong stomatal control but high photosynthetic rates, which increases water use efficiency (iWUE), and an enhanced ability to accumulate sugars as osmolytes. Although drought decreased RGR in all populations, this reduction was smaller for populations from the dry edge. Our results suggest that dry-edge populations of this relict species are well adapted to drought, which could potentially mitigate the species' extinction risk under drier scenarios. Dry-edge populations not only have a great conservation value but can also change expectations from current species' distribution models.
Extensive within-canopy light gradients importantly affect the photosynthetic productivity of leaves in different canopy positions and lead to light-dependent increases in foliage photosynthetic capacity per area (AA). However, the controls on AA variations by changes in underlying traits are poorly known. We constructed an unprecedented worldwide database including 831 within-canopy gradients with standardized light estimates for 304 species belonging to major vascular plant functional types, and analyzed within-canopy variations in 12 key foliage structural, chemical and physiological traits by quantitative separation of the contributions of different traits to photosynthetic acclimation. Although the light-dependent increase in AA is surprisingly similar in different plant functional types, they differ fundamentally in the share of the controls on AA by constituent traits. Species with high rates of canopy development and leaf turnover, exhibiting highly dynamic light environments, actively change AA by nitrogen reallocation among and partitioning within leaves. By contrast, species with slow leaf turnover exhibit a passive AA acclimation response, primarily determined by the acclimation of leaf structure to growth light. This review emphasizes that different combinations of traits are responsible for within-canopy photosynthetic acclimation in different plant functional types, and solves an old enigma of the role of mass- vs area-based traits in vegetation acclimation.
To interpret time series of hyperspectral measurements of vegetation canopies, the basic characteristics of the scattering elements forming these canopies-shoots, leaves, or needles-must be known. Unfortunately, data on the seasonal variation of leaf reflectance and transmittance are very scarce. To obtain a ground truth dataset applicable to modeling the phenological development of European (hemi) boreal forests, we measured the spectral properties of green leaves of two species common to this biome, gray alder (Alnus incana), and silver birch (Betula pendula). Measurements covered the full growing season of 2008 in Toravere, Estonia. Leaves were sampled from sunlit locations and measured in a laboratory using an integrating sphere and a VNIR spectroradiometer. We measured four different optical parameters: directional-hemispherical reflectance and transmittance factors for leaf adaxial and abaxial surfaces. Leaf reflectance was used to calculate four leaf-level indices which, according to literature, are highly correlated with leaf chlorophyll content. Additionally, we calculated the derivative spectra of leaf reflectance in the red edge (RE) spectral region and fitted it with two Gaussian curves. Our analysis indicated continuous changes of leaf optical properties almost until the end of the growing season. The changes in the weights of the two Gaussian curves led to an effect known as peak jump, an abrupt shift in the RE inflection point. In near infrared (NIR), leaf absorption was negligible in the beginning of the growing period. However, we noted a slow but steady increase in the leaf NIR absorption with time.
Foliage photosynthetic and structural traits were studied in 15 species with a wide range of foliage anatomies to gain insight into the importance of key anatomical traits in the limitation of diffusion of CO2 from substomatal cavities to chloroplasts. The relative importance of different anatomical traits in constraining CO2 diffusion was evaluated using a quantitative model. Mesophyll conductance (gm) was most strongly correlated with chloroplast exposed surface to leaf area ratio (Sc/S) and cell wall thickness (Tcw), but, depending on foliage structure, the overall importance of gm in constraining photosynthesis and the importance of different anatomical traits in the restriction of CO2 diffusion varied. In species with mesophytic leaves, membrane permeabilities and cytosol and stromal conductance dominated the variation in gm. However, in species with sclerophytic leaves, gm was mostly limited by Tcw. These results demonstrate the major role of anatomy in constraining mesophyll diffusion conductance and, consequently, in determining the variability in photosynthetic capacity among species.
INCREASE infrastructures (EU 7th FP) provide access for visiting scientists to study the effects of long-term climate manipulation treatments in field experiments on shrubland ecosystems across Europe. Climate manipulations include prolonged drought and warming treatments which have lasted for over 10 years, and free-air increased CO2 treatment in Denmark, Brandbjerg running over 5 years. Non-invasive measurement techniques are particularly encouraged for minimizing the disturbance on experimental plots. We visited the experimental sites in Italy (Porto Conte) and Denmark (Mols and Brandbjerg) in summer 2011 to conduct reflectance measurements in visible and short-wave infrared spectral region (400-2500 nm).
We investigated changes in chlorophyll a fluorescence from alternate leaf surfaces to assess the intraleaf light acclimation patterns in combination with natural variations in radiation, leaf angles, leaf mass per area (LMA), chlorophyll content (Chl) and leaf optical parameters. Measurements were conducted on bottom- and top-layer leaves of Tilia cordata Mill. (a shade-tolerant sub-canopy species, sampled at heights of 11 and 16 m) and Populus tremula L. (a light-demanding upper canopy species, sampled at canopy heights of 19 and 26 m). The upper canopy species P. tremula had a six times higher PSII quantum yield (Φ(II)) and ratio of open reaction centres (qP), and a two times higher LMA than T. cordata. These species-specific differences were also present when the leaves of both species were in similar light conditions. Leaf adaxial/abaxial fluorescence ratio was significantly larger in the case of more horizontal leaves. Populus tremula (more vertical leaves), had smaller differences in fluorescence parameters between alternate leaf sides compared with T. cordata (more horizontal leaves). However, optical properties on alternate leaf sides showed a larger difference for P. tremula. Intraspecifically, the measured optical parameters were better correlated with LMA than with leaf Chl. Species-specific differences in leaf anatomy appear to enhance the photosynthetic potential of leaf biochemistry by decreasing the interception of excess light in P. tremula and increasing the light absorptance in T. cordata. Our results indicate that intraleaf light absorption gradient, described here as leaf adaxial/abaxial side ratio of chlorophyll a fluorescence, varies significantly with changes in leaf light environment in a multi-layer multi-species tree canopy. However, this variation cannot be described merely as a simple function of radiation, leaf angle, Chl or LMA, and species-specific differences in light acclimation strategies should also be considered.
Needle morphological, chemical and physiological characteristics of Norway spruce were studied in a forest chronosequence in Järvselja Experimental Forest, Estonia. Current-year shoots were sampled from upper canopy positions in five stands, ranging in height from 1.8 to 33.0 m (corresponding age range was 10-85 years). A/C(i) curves were determined to obtain maximum carboxylation rates (V(cmax)) and maximum rates of electron transport (J(max)). Needle nitrogen (N) partitioning into photosynthetic functions was calculated from the values of V(cmax), J(max) and leaf chlorophyll concentration. All needle size parameters (length, width, thickness, volume and cross-sectional areas of mesophyll and xylem) increased significantly with tree height. The needles of taller trees had lower mass-based N and chlorophyll concentrations (21% and 43% difference between shortest and tallest stands, respectively), but higher dry mass per area (35%), dry mass per volume (18%), number of cells per mesophyll cross-section area (40%) and partitioning of N into non-photosynthetic functions (12%). Light saturated net assimilation rate, V(cmax), J(max) and stomatal conductance decreased with tree age (35%, 16%, 12% and 29% difference, respectively). A path analysis model describing tree age-related reduction of photosynthetic capacity as a result of sink limitation provided the best fit to our data. However, since the path model corresponding to source limitation, where photosynthetic reduction derives from changes in needle structure and chemistry was not rejected, we conclude that the decline in photosynthesis with tree age results from several mechanisms (limited sink strength, stomatal and N limitation) operating simultaneously and sequentially.
Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.
The aim of this study was to evaluate the use of ground-based canopy reflectance measurements to detect changes in physiology and structure of vegetation in response to experimental warming and drought treatment at six European shrublands located along a North–South climatic gradient. We measured canopy reflectance, effective green leaf area index (green LAIe) and chlorophyll fluorescence of dominant species. The treatment effects on green LAIe varied among sites. We calculated three reflectance indices: photochemical reflectance index PRI [531nm; 570nm], normalized difference vegetation index NDVI680 [780nm; 680nm] using red spectral region, and NDVI570 [780nm; 570nm] using the same green spectral region as PRI. All three reflectance indices were significantly related to green LAIe and were able to detect changes in shrubland vegetation among treatments. In general warming treatment increased PRI and drought treatment reduced NDVI values. The significant treatment effect on photochemical efficiency of plants detected with PRI could not be detected by fluorescence measurements. However, we found canopy level measured PRI to be very sensitive to soil reflectance properties especially in vegetation areas with low green LAIe. As both soil reflectance and LAI varied between northern and southern sites it is problematic to draw universal conclusions of climate-derived changes in all vegetation types based merely on PRI measurements. We propose that canopy level PRI measurements can be more useful in areas of dense vegetation and dark soils.
The reflectance signal from the forest tree canopy is influenced by the optical properties of the background formed by understory vegetation. Our study shows that the herb-moss layer in the forest tends to be brighter in the visible wavelength when the canopy above is more closed because of the specific properties of plants grown in low light. When leaf-area-based chlorophyll content falls below approximately 150 mg·m–2, reflectance in the red region of the spectrum increases compared with that of the background. The best descriptors of the herb-moss layer for deriving optical parameters are herb layer dry mass for the visible wavelength range and total aboveground water mass for near-infrared (NIR) reflectance. In addition, chlorophyll content per leaf area considerably improves the red reflectance estimate.
Leaf-level determinants of species environmental stress tolerance are still poorly understood. Here, we explored dependencies of species shade (T(shade)) and drought (T(drought)) tolerance scores on key leaf structural and functional traits in 339 Northern Hemisphere temperate woody species. In general, T(shade) was positively associated with leaf life-span (L(L)), and negatively with leaf dry mass (M(A)), nitrogen content (N(A)), and photosynthetic capacity (A(A)) per area, while opposite relationships were observed with drought tolerance. Different trait combinations responsible for T(shade) and T(drought) were observed among the key plant functional types: deciduous and evergreen broadleaves and evergreen conifers. According to principal component analysis, resource-conserving species with low N content and photosynthetic capacity, and high L(L) and M(A), had higher T(drought), consistent with the general stress tolerance strategy, whereas variation in T(shade) did not concur with the postulated stress tolerance strategy. As drought and shade often interact in natural communities, reverse effects of foliar traits on these key environmental stress tolerances demonstrate that species niche differentiation is inherently constrained in temperate woody species. Different combinations of traits among key plant functional types further explain the contrasting bivariate correlations often observed in studies seeking functional explanation of variation in species environmental tolerances.
We studied acclimation patterns in leaf dry mass per area (MA), nitrogen (NA) and chlorophyll (ζA) content per area, and chlorophyll to nitrogen ratio (ζ/N) along vertical light gradients in natural temperate mixed herbaceous canopy and deciduous tree canopy. In the deciduous tree canopy, all leaves are formed at approximately the same time, and the light gradient during the rest of the growing season reflects the differences in light availability during leaf development, whereas in the herbaceous canopy, leaf production continues during most of the growing season and major changes in light conditions occur after leaf maturation. MA and NA increased strongly with increasing current light availability (ID) in the tree canopy. In the herbaceous canopy, MA and NA were generally unrelated to ID. Depending on species, the correlation between chlorophyll content per leaf area (ζA) and ID was positive, negative, or non-significant. Path analyses revealed two opposite effects of ID on the amount of leaf chlorophyll. In the tree canopy, increasing ID enhanced ζA through changes in MA and NA, whereas the direct effect of light was negative in both canopies. The overall correlation network between foliage structural and chemical traits and the relationships with ID were significantly stronger in the tree canopy, suggesting limited re-acclimation potential in the mixed herbaceous canopy. Within-species acclimation patterns reflected the patterns within the main functional types. These data demonstrate that the relationships of current light availability vs. leaf dry mass per area, leaf nitrogen and chlorophyll contents, and chlorophyll to nitrogen ratio differ among multi-species herbaceous canopies and deciduous tree canopies due to contrasting canopy development.