Unravelling the neglected role of ultraviolet radiation on stomata: A meta-analysis with implications for modelling ecosystem-climate interactions.

Stomata play a pivotal role in regulating gas exchange between plants and the atmosphere controlling water and carbon cycles. Accordingly, we investigated the impact of ultraviolet-B radiation, a neglected environmental factor varying with ongoing global change, on stomatal morphology and function by a Comprehensive Meta-Analysis. The overall UV effect at the leaf level is to decrease stomatal conductance, stomatal aperture and stomatal size, although stomatal density was increased. The significant decline in stomatal conductance is marked (6% in trees and >10% in grasses and herbs) in short-term experiments, with more modest decreases noted in long-term UV studies. Short-term experiments in growth chambers are not representative of long-term field UV effects on stomatal conductance. Important consequences of altered stomatal function are hypothesized. In the short term, UV-mediated stomatal closure may reduce carbon uptake but also water loss through transpiration, thereby alleviating deleterious effects of drought. However, in the long term, complex changes in stomatal aperture, size, and density may reduce the carbon sequestration capacity of plants and increase vegetation and land surface temperatures, potentially exacerbating negative effects of drought and/or heatwaves. Therefore, the expected future strength of carbon sink capacity in high-UV regions is likely overestimated.

A meta-analysis of the interactive effects of UV and drought on plants.

Interactions between climate change and UV penetration in the biosphere are resulting in the exposure of plants to new combinations of UV radiation and drought. In theory, the impacts of combinations of UV and drought may be additive, synergistic or antagonistic. Lack of understanding of the impacts of combined treatments creates substantial uncertainties that hamper predictions of future ecological change. Here, we compiled information from 52 publications and analysed the relative impacts of UV and/or drought. Both UV and drought have substantial negative effects on biomass accumulation, plant height, photosynthesis, leaf area and stomatal conductance and transpiration, while increasing stress-associated symptoms such as MDA accumulation and reactive-oxygen-species content. Contents of proline, flavonoids, antioxidants and anthocyanins, associated with plant acclimation, are upregulated both under enhanced UV and drought. In plants exposed to both UV and drought, increases in plant defense responses are less-than-additive, and so are the damage and growth retardation. Less-than-additive effects were observed across field, glasshouse and growth-chamber studies, indicating similar physiological response mechanisms. Induction of a degree of cross-resistance seems the most likely interpretation of the observed less-than-additive responses. The data show that in future climates, the impacts of increases in drought exposure may be lessened by naturally high UV regimes.

Response of green reflectance continuum removal index to the xanthophyll de-epoxidation cycle in Norway spruce needles.

A dedicated field experiment was conducted to investigate the response of a green reflectance continuum removal-based optical index, called area under the curve normalized to maximal band depth between 511 nm and 557 nm (ANMB511-557), to light-induced transformations in xanthophyll cycle pigments of Norway spruce [Picea abies (L.) Karst] needles. The performance of ANMB511-557 was compared with the photochemical reflectance index (PRI) computed from the same leaf reflectance measurements. Needles of four crown whorls (fifth, eighth, 10th, and 15th counted from the top) were sampled from a 27-year-old spruce tree throughout a cloudy and a sunny day. Needle optical properties were measured together with the composition of the photosynthetic pigments to investigate their influence on both optical indices. Analyses of pigments showed that the needles of the examined whorls varied significantly in chlorophyll content and also in related pigment characteristics, such as the chlorophyll/carotenoid ratio. The investigation of the ANMB511-557 diurnal behaviour revealed that the index is able to follow the dynamic changes in the xanthophyll cycle independently of the actual content of foliar pigments. Nevertheless, ANMB511-557 lost the ability to predict the xanthophyll cycle behaviour during noon on the sunny day, when the needles were exposed to irradiance exceeding 1000 µmol m(-2) s(-1). Despite this, ANMB511-557 rendered a better performance for tracking xanthophyll cycle reactions than PRI. Although declining PRI values generally responded to excessive solar irradiance, they were not able to predict the actual de-epoxidation state in the needles examined.

Estimation of photosynthetic dynamics in forests from daily measured fluorescence and PRI data with adjustment for canopy shadow fraction.

We conducted year-long measurements of the photochemical reflectance index (PRI) and solar-induced fluorescence in the O2A oxygen band (SIFA) at a Norway spruce forest and a European beech forest to study relationships of these remote sensing variables to photosynthesis by trees in grown forest stands. Measured PRI and SIFA values were linked to changes in forest gross primary productivity (GPP) and light-use efficiency (LUE). Changes in the shadow fraction (αS) within tree crowns influenced PRI and fluorescence signals. In the spruce forest, the quantum yield of SIFA (FYSIFA) decreased around midday together with photosynthesis and GPP. Such decreases in FYSIFA were accompanied by an increase in the αS. In the beech forest, we detected an increase in FYSIFA together with a decrease in αS in the afternoon hours. The overall sensitivity of PRI to LUE was variable according to the season, presumably influenced by complex changes in photosynthetic pigments. PRI and FYSIFA showed weak correlations with canopy LUE; however, when considered together, the correlation was strengthened (R2 was 0.63 and 0.34 in spruce and beech forest, respectively). Our model predicting LUE dynamics includes a diurnal minimum of PRI and canopy αS to make allowances for seasonal changes in photosynthetic pigments and for diurnal variability of the shadow fraction in forests. The incorporation of these correcting factors allowed us to estimate LUE at R2 = 0.68 (spruce) and 0.53 (beech). The modeling equations appeared sensitive to the absorbed photosynthetically active radiation (APAR), but less sensitive to the GPP of these forests. Substituting pigments correction with introducing differential PRI (ΔPRI) into the model did not significantly improve the LUE estimation across the season. Our results show that the joint use of PRI and fluorescence improves LUE and GPP estimation accuracy in both daily and seasonal observations.

Relation of chlorophyll fluorescence sensitive reflectance ratios to carbon flux measurements of montanne grassland and norway spruce forest ecosystems in the temperate zone.

We explored ability of reflectance vegetation indexes (VIs) related to chlorophyll fluorescence emission (R686/R630, R740/R800) and de-epoxidation state of xanthophyll cycle pigments (PRI, calculated as (R531- R570)/(R531-R570) to track changes in the CO2 assimilation rate and Light Use Efficiency (LUE) in montane grassland and Norway spruce forest ecosystems, both at leaf and also canopy level. VIs were measured at two research plots using a ground-based high spatial/spectral resolution imaging spectroscopy technique. No significant relationship between VIs and leaf light-saturated CO2 assimilation (A(MAX)) was detected in instantaneous measurements of grassland under steady-state irradiance conditions. Once the temporal dimension and daily irradiance variation were included into the experimental setup, statistically significant changes in VIs related to tested physiological parameters were revealed. ΔPRI and Δ(R686/R630) of grassland plant leaves under dark-to-full sunlight transition in the scale of minutes were significantly related to A(MAX) (R² = 0.51). In the daily course, the variation of VIs measured in one-hour intervals correlated well with the variation of Gross Primary Production (GPP), Net Ecosystem Exchange (NEE), and LUE estimated via the eddy-covariance flux tower. Statistical results were weaker in the case of the grassland ecosystem, with the strongest statistical relation of the index R686/R630 with NEE and GPP.

A meta-analysis of the effects of UV radiation on the plant carotenoid pool.

Induction of metabolite biosynthesis and accumulation is one of the most prominent UV-mediated changes in plants, whether during eustress (positive response) or distress (negative response). However, despite evidence suggesting multiple linkages between UV exposure and carotenoid induction in plants, there is no consensus in the literature concerning the direction and/or amplitude of these effects. Here, we compiled publications that characterised the relative impact of UV on the content of individual carotenoids and subjected the created database to a meta-analysis in order to acquire new, fundamental insights in responses of the carotenoid pool to UV exposure. Overall, it was found that violaxanthin was the only carotenoid compound that was significantly and consistently induced as a result of UV exposure. Violaxanthin accumulation was accompanied by a UV dose dependent decrease in antheraxanthin and zeaxanthin. The resulting shift in the state of the xanthophyll cycle would normally occur when plants are exposed to low light and this is associated with increased susceptibility to photoinhibition. Although UV induced violaxanthin accumulation is positively linked to the daily UV dose, the current dataset is too small to establish a link with plant stress, or even experimental growth conditions. In summary, the effects of UV radiation on carotenoids are multifaceted and compound-specific, and there is a need for a systematic analysis of dose-response and wavelength dependencies, as well as of interactive effects with further environmental parameters.

Combining NDVI, PRI and the quantum yield of solar-induced fluorescence improves estimations of carbon fluxes in deciduous and evergreen forests.

We used automated spectroradiometers to continuously monitor changes in the optical parameters of phenological and photosynthetic traits in beech and spruce forests. We examined seasonal variations in the normalized difference vegetation index (NDVI), photochemical reflectance index (PRI), and solar-induced fluorescence in the oxygen A band (SIFA) that was estimated using a 3-FLD discrimination method from radiance data. The optical parameters tracked the activation and cessation of photosynthesis in spring and autumn. Data at photon fluxes >1200 µmol m-2 s-1 during extended noon hours were used to link the seasonal PRI and SIFA variations to the dynamics of photosynthesis. Seasonal PRI was significantly correlated with photosynthetic light-use efficiency (LUE) with R2 values of 0.66 and 0.48 for the measurements in beech and spruce forests, respectively. SIFA emissions were significantly correlated with the gross primary production (GPP) of the evergreen spruce forest (R2 = 0.47), but R2 was only 0.13 when measured in the beech forest. The correlations between the optical parameters and GPP or LUE, however, tended to be lower when using a dataset with constant NDVI. Introducing an equation combining NDVI, PRI, and the quantum yield of SIFA emission increased R2 for LUE estimation to 0.77 in the spruce forest and 0.63 in the beech forest. GPP was estimated from the parametric equation with improved accuracy reaching R2 = 0.53 and RMSE = 5.95 µmol CO2 m-2 s-1 in spruce forest and R2 = 0.58 and RMSE = 5.23 µmol CO2 m-2 s-1 in beech forest. Parametric equations were more efficient in estimating photosynthesis in datasets that consisted of an entire season's data. By combining NDVI, PRI and the quantum yield of SIFA, we could thus substantially improve estimations of carbon fluxes in diverse deciduous and evergreen canopies.

Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species.

The differential pigment composition and photosynthetic activity of sun and shade leaves of deciduous (Acer pseudoplatanus, Fagus sylvatica, Tilia cordata) and coniferous (Abies alba) trees was comparatively determined by studying the photosynthetic rates via CO(2) measurements and also by imaging the Chl fluorescence decrease ratio (R(Fd)), which is an in vivo indicator of the net CO(2) assimilation rates. The thicker sun leaves and needles in all tree species were characterized by a lower specific leaf area, lower water content, higher total chlorophyll (Chl) a+b and total carotenoid (Cars) content per leaf area unit, as well as higher values for the ratio Chl a/b compared to the much thinner shade leaves and needles that possess a higher Chl a+b and Cars content on a dry matter basis and higher values for the weight ratio Chls/Cars. Sun leaves and needles exhibited higher rates of maximum net photosynthetic CO(2) assimilation (P(Nmax)) measured at saturating irradiance associated with higher maximum stomatal conductance for water vapor efflux. The differences in photosynthetic activity between sun and shade leaves and needles could also be sensed via imaging the Chl fluorescence decrease ratio R(Fd), since it linearly correlated to the P(Nmax) rates at saturating irradiance. Chl fluorescence imaging not only provided the possibility to screen the differences in P(N) rates between sun and shade leaves, but in addition permitted detection and quantification of the large gradients in photosynthetic rates across the leaf area existing in sun and shade leaves.

Near-distance imaging spectroscopy investigating chlorophyll fluorescence and photosynthetic activity of grassland in the daily course.

Detection of grassland canopy chlorophyll fluorescence (Chl-F) conducted with an imaging spectroradiometer provided evidence of potential remote sensing estimation of steady-state Chl-F (Chl-Fs). Daily near-nadir views of extremely high spatial resolution hyperspectral images were acquired from a distance of 4 m for temperate montane grassland in the Czech Republic. Simultaneously, measurements of Chl-F and total chlorophyll content (Chla + b) were made on a single leaf at ground level were collected. A specifically designed 'shade removal' experiment revealed the influence of dynamic physiological plant processes on hyperspectral reflectance of three wavelengths: 532, 686 and 740 nm. Based on this information, the vegetation indexes R686/R630, R740/R800 and PRI calculated as (R532-R570)/(R532+R570) were tested for statistical significance with directly measured Chl-F parameters (maximum fluorescence yield, Fv/Fm; steady-state chlorophyll fluorescence, Chl-Fs and actual quantum yield, ФII). The grassland species under investigation were: Festuca rubra agg. (L.), Hieracium sp., Plantago sp., Nardus stricta (L.) and Jacea pseudophrygia (C.A. Meyer). The coefficients of determination (R2) for best-fit relationships between PRI-ФII and PRI-Chl-Fs, measured in the daily course, show a high variability of 0.23-0.78 and 0.20-0.65, respectively. Similarly, R2 for the R686/R630-ФII and R686/R630-Chl-Fs relationships varied between 0.20-0.73 and 0.41-0.70, respectively. The highest average R2 values were found between PRI and Chla + b (0.63) and R686/R630 and Chla + b (0.72). The ratio R740/R800 did not yield a statistically significant relation with Chl-F parameters.