Relationship between photochemical reflectance index and leaf ecophysiological and biochemical parameters under two different water statuses: towards a rapid and efficient correction method using real-time measurements.

The use of the photochemical reflectance index (PRI) as a promising proxy of light use efficiency (LUE) has been extensively studied, and some issues have been identified, notably the sensitivity of PRI to leaf pigment composition and the variability in PRI response to LUE because of stress. In this study, we introduce a method that enables us to track the short-term PRI response to LUE changes because of photosynthetically active radiation (PAR) changes. The analysis of these short-term relationships between PRI and LUE throughout the growing season in two species (Quercus robur L. and Fagus sylvatica L.) under two different soil water statuses showed a clear change in PRI response to LUE, which is related to leaf pigment content. The use of an estimated or approximated PRI0, defined as the PRI of perfectly dark-adapted leaves, allowed us to separate the PRI variability due to leaf pigment content changes and the physiologically related PRI variability over both daily (PAR-related) and seasonal (soil water content-related) scales. The corrected PRI obtained by subtracting PRI0 from the PRI measurements showed a good correlation with the LUE over both of the species, soil water statuses and over the entire growing season.

Light and competition gradients fail to explain the coexistence of shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings.

BACKGROUND AND AIMS: The coexistence of forest tree species has often been linked to differences among species in terms of their response to light availability during the regeneration stage. From this perspective, species coexistence results from growth-growth or mortality-growth trade-offs along spatial light gradients. Experimental evidence of growth-growth trade-offs in natural conditions is sparse due to various confounding factors that potentially hinder the relationship. This study examined growth hierarchies along light gradients between two tree species with contrasting shade tolerance by controlling potential confounding factors such as seedling size, seedling status, seedling density and species composition. METHODS: Natural regenerated shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings were used, and growth rankings over a 4-year period were compared in 8- to 10-year-old tree seedlings. KEY RESULTS: No rank reversal occurs between the two species along the light gradient, or along the density, mixture or seedling size gradients. The shade-tolerant species was always the more competitive of the two. Pronounced effects of initial size on seedling growth were observed, whereas the effects of light and competition by neighbours were of secondary importance. The paramount effect of size, which results from the asymmetric nature of interseedling competition, gives a strong advantage to tall seedlings over the long term. CONCLUSIONS: This study extends previous efforts to identify potential drivers of rank reversals in young tree mixtures. It does not support the classical assumption that spatial heterogeneity in canopy opening explains the coexistence of the two species studied. It suggests that spatial variation in local size hierarchies among seedlings that may be caused by seedling emergence time or seedling initial performance is the main driver of the dynamics of these mixed stands.

Growth and posture control strategies in Fagus sylvatica and Acer pseudoplatanus saplings in response to canopy disturbance.

BACKGROUND AND AIMS: Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation. METHODS: A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening. KEY RESULTS: Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth. CONCLUSIONS: Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.

Insights into xylem vulnerability to cavitation in Fagus sylvatica L.: phenotypic and environmental sources of variability.

Xylem vulnerability to cavitation is a key parameter in understanding drought resistance of trees. We determined the xylem water pressure causing 50% loss of hydraulic conductivity (P(50)), a proxy of vulnerability to cavitation, and we evaluated the variability of this trait at tree and population levels for Fagus sylvatica. We checked for the effects of light on vulnerability to cavitation of stem segments together with a time series variation of P(50). Full sunlight-exposed stem segments were less vulnerable to cavitation than shade-exposed ones. We found no clear seasonal change of P(50), suggesting that this trait was designed for a restricted period. P(50) varied for populations settled along a latitudinal gradient, but not for those sampled along an altitudinal gradient. Moreover, mountainside exposure seemed to play a major role in the vulnerability to cavitation of beech populations, as we observed the differences along north-facing sides but not on south-facing sides. Unexpectedly, both north-facing mountainside and northern populations appeared less vulnerable than those grown on the southern mountainside or in the South of France. These results on beech populations were discussed with respect to the results at within-tree level.

Gregarious bamboo flowering opens a window of opportunity for regeneration in a temperate forest of Patagonia.

Rare gregarious flowering of understorey bamboo species occurs in temperate and subtropical forests around the world, but the ecological consequences of this phenomenon for forest regeneration are not well understood.Field experiments were conducted in an old-growth temperate forest in Patagonia,Argentina after a massive bamboo flowering event, to examine whether light quality and other changes in microhabitats could affect seed germination and growth of overstorey species. Germination of southern beech (Nothofagus obliqua) was positively correlated with red:far red (R:FR) ratios in a range of microhabitats generated by the death of the understorey bamboo (Chusquea culeou). Experimental modification of understorey R:FR ratios to mimic alternative light environments reversed this germination response in plots with senescent understorey, but not in plots with live bamboo. Laboratory incubations demonstrated a significant interaction between R:FR ratios and thermal amplitude in promoting seed germination. Microhabitats also significantly affected the growth of emerged seedlings. Microenvironmental changes generated by this flowering event appear to have opened a window of opportunity for germination and growth of overstorey species.We demonstrate that natural gradients in light quality associated with this ecological phenomenon are a major component affecting forest regeneration in this ecosystem.

Seasonal time-course of gradients of photosynthetic capacity and mesophyll conductance to CO2 across a beech (Fagus sylvatica L.) canopy.

Leaf photosynthesis is known to acclimate to the actual irradiance received by the different layers of a canopy. This acclimation is usually described in terms of changes in leaf structure, and in photosynthetic capacity. Photosynthetic capacity is likely to be affected by mesophyll conductance to CO(2) which has seldom been assessed in tree species, and whose plasticity in response to local irradiance is still poorly known. Structural [N and chlorophyll content, leaf mass to area ratio (LMA)] and functional leaf traits [maximum carboxylation rate (V(cmax)), maximum light-driven electron flux (J(max)), and mesophyll conductance (g(i))] were assessed by measuring leaf response curves of net CO(2) assimilation versus intercellular CO(2) partial pressure, along a vertical profile across a beech canopy, and by fitting a version of the Farquhar model including g(i). The measurements were repeated five times during a growth season to catch potential seasonal variation. Irradiance gradients resulted in large decreasing gradients of LMA, g(i), V(cmax), and J(max). Relative allocation of leaf N to the different photosynthetic processes was only slightly affected by local irradiance. Seasonal changes after leaf expansion and before induction of leaf senescence were only minor. Structural equation modelling confirmed that LMA was the main driving force for changes in photosynthetic traits, with only a minor contribution of leaf Nitrogen content. In conclusion, mesophyll conductance to CO(2) displays a large plasticity that scales with photosynthetic capacity across a tree canopy, and that it is only moderately (if at all) affected by seasonal changes in the absence of significant soil water depletion.

Within-canopy and ozone fumigation effects on delta13C and Delta18O in adult beech (Fagus sylvatica) trees: relation to meteorological and gas exchange parameters.

In this study, the effects of different light intensities either in direct sunlight or in the shade crown of adult beech (Fagus sylvatica L.) trees on delta13C and Delta18O were determined under ambient (1 x O3) and twice-ambient (2 x O3) atmospheric ozone concentrations during two consecutive years (2003 and 2004). We analysed the isotopic composition in leaf bulk, leaf cellulose, phloem and xylem material and related the results to (a) meteorological data (air temperature, T and relative humidity, RH), (b) leaf gas exchange measurements (stomatal conductance, g(s); transpiration rate, E; and maximum photosynthetic activity, A(max)) and (c) the outcome of a steady-state evaporative enrichment model. Delta13C was significantly lower in the shade than in the sun crown in all plant materials, whilst Delta18O was increased significantly in the shade than in the sun crown in bulk material and cellulose. Elevated ozone had no effect on delta13C, although Delta18O was influenced by ozone to varied degrees during single months. We observed significant seasonal changes for both parameters, especially in 2004, and also significant differences between the study years. Relating the findings to meteorological data and gas exchange parameters, we conclude that the differences in Delta18O between the sun and the shade crown were predominantly caused by the Péclet effect. This assumption was supported by the modelled Delta18O values for leaf cellulose. It was demonstrated that independent of RH, light-dependent reduction of stomatal conductance (and thus transpiration) and of A(max) can drive the pattern of Delta18O increase with the concomitant decrease of delta13C in the shade crown. The effect of doubling ozone levels on time-integrated stomatal conductance and transpiration as indicated by the combined analysis of Delta18O and delta13C was much lower than the influence caused by the light exposure.

Ultraviolet radiation modulates C:N stoichiometry and biomass allocation in Fagus sylvatica saplings cultivated under elevated CO2 concentration.

Under the conditions of ongoing climate change, terrestrial ecosystems will be simultaneously exposed to a permanent rise in atmospheric CO2 concentration and increasing variability of such environmental factors as temperature, precipitation, and UV radiation. This will result in numerous interactions. The interactive effects caused by exposure to such multiple environmental factors are not yet well understood. We tested the hypotheses that enhanced UV radiation reduces the stimulatory effect of elevated CO2 concentration on plant biomass production and that it alters biomass allocation in broadleaved European beech (Fagus sylvatica L.) saplings. Our results after 2 years of exposure confirmed interactive effects of CO2 concentration and UV radiation on biomass production, and particularly on biomass allocation to roots and aboveground biomass. The strongest stimulatory effect of elevated CO2 on aboveground biomass and roots was found under ambient UV radiation, while both low and high UV doses reduced this stimulation. Nitrogen content in the roots and the distribution of nitrogen among leaves and roots were also significantly affected by interaction of CO2 concentration and UV radiation. The observed changes in leaf and root C:N stoichiometry were associated with altered morphological traits, and particularly with a change in the proportion of fine roots. As the biomass allocation and especially the proportion of fine roots can play an important role in effective water and nutrient use and acclimation to future climates, it is essential to obtain a deeper understanding of the links between C:N stoichiometry and biomass accumulation.

Interactive effects of ultraviolet radiation and elevated CO2 concentration on photosynthetic characteristics of European beech saplings during the vegetation season.

To test the hypothesis that ultraviolet radiation (UV) modulates photosynthetic responses to elevated CO2 concentration ([CO2]) in plants, saplings of European beech were grown for two vegetation seasons under ambient (400 ppm) and elevated (700 ppm) atmospheric [CO2]. From April to November the saplings were exposed to (i) ambient UV radiation, (ii) excluded and (iii) enhanced UV (150% of ambient). Gas-exchange and chlorophyll fluorescence techniques were used throughout the second vegetation season together with biochemical analyses of the amount and activity of the Rubisco enzyme. We found support for the hypothesis that an impact of elevated [CO2] on photosynthesis is substantially modulated by UV radiation. Moreover, we found that the [CO2] × UV interaction is changing along the vegetation season: an enhanced UV radiation stimulated a positive effect of elevated [CO2] on plant photosynthesis at the beginning of the vegetation season (short-term effect), whilst long-term cultivation reduced the stimulatory effect of elevated [CO2] (a clear down-regulation of photosynthesis). Down-regulation was, however, not found in plants grown under the conditions of excluded UV radiation. We found evidence that the down-regulation of photosynthesis is associated with a complex acclimation at different hierarchical and functional levels, including an acclimation of primary photochemical reactions, carboxylation activity of Rubisco enzyme, and stomatal conductance.

Ontogenetic variation in the relative influence of light and belowground resources on European beech seedling growth.

We used height growth data from a 7-year field experiment with European beech (Fagus sylvatica L.) seedlings to test the hypothesis that the effects of above- and belowground resources on height growth depend on seedling size and age. Photosynthetically active radiation (PAR) was determined by hemispheric photography, and estimates of fine root biomass of the overstory trees were used as an inverse proportional surrogate for belowground resource availability. For recently germinated seedlings growing under the canopy of Picea abies (L.) Karst., belowground resource availability affected height growth more than light. During subsequent stages of seedling development, apart from initial seedling size, PAR increasingly determined seedling growth. Besides initial size, seedling age determined the effects of above- and belowground resources on seedling height growth. In seedlings identical in initial size but differing in age, the increase in height growth with increasing PAR was greater in older seedlings than in younger seedlings. The ranking of seedling height by year showed that small differences in size at the end of the first growing season resulted in continuously increasing differences during the following years. Mortality data indicated that the chances of a seedling surviving intraspecific competition was strongly determined by its dominance ranking within the first 5 years after establishment.

CASIROZ: Root parameters and types of ectomycorrhiza of young beech plants exposed to different ozone and light regimes.

Tropospheric ozone (O(3)) triggers physiological changes in leaves that affect carbon source strength leading to decreased carbon allocation below-ground, thus affecting roots and root symbionts. The effects of O(3) depend on the maturity-related physiological state of the plant, therefore adult and young forest trees might react differently. To test the applicability of young beech plants for studying the effects of O(3) on forest trees and forest stands, beech seedlings were planted in containers and exposed for two years in the Kranzberg forest FACOS experiment (Free-Air Canopy O(3) Exposure System, http://www.casiroz.de ) to enhanced ozone concentration regime (ambient [control] and double ambient concentration, not exceeding 150 ppb) under different light conditions (sun and shade). After two growing seasons the biomass of the above- and below-ground parts, beech roots (using WinRhizo programme), anatomical and molecular (ITS-RFLP and sequencing) identification of ectomycorrhizal types and nutrient concentrations were assessed. The mycorrhization of beech seedlings was very low ( CA. 5 % in shade, 10 % in sun-grown plants), no trends were observed in mycorrhization (%) due to ozone treatment. The number of Cenococcum geophilum type of ectomycorrhiza, as an indicator of stress in the forest stands, was not significantly different under different ozone treatments. It was predominantly occurring in sun-exposed plants, while its majority share was replaced by Genea hispidula in shade-grown plants. Different light regimes significantly influenced all parameters except shoot/root ratio and number of ectomycorrhizal types. In the ozone fumigated plants the number of types, number of root tips per length of 1 to 2 mm root diameter, root length density per volume of soil and concentration of Mg were significantly lower than in control plants. Trends to a decrease were found in root, shoot, leaf, and total dry weights, total number of root tips, number of vital mycorrhizal root tips, fine root (mass) density, root tip density per surface, root area index, concentration of Zn, and Ca/Al ratio. Due to the general reduction in root growth indices and nutrient cycling in ozone-fumigated plants, alterations in soil carbon pools could be predicted.

O3 flux-related responsiveness of photosynthesis, respiration, and stomatal conductance of adult Fagus sylvatica to experimentally enhanced free-air O3 exposure.

Knowledge of responses of photosynthesis, respiration, and stomatal conductance to cumulative ozone uptake (COU) is still scarce, and this is particularly the case for adult trees. The effect of ozone (O(3)) exposure on trees was examined with 60-year-old beech trees (FAGUS SYLVATICA) at a forest site of southern Germany. Trees were exposed to the ambient O(3) regime (1 x O(3)) or an experimentally elevated twice-ambient O(3) regime (2 x O(3)). The elevated 2 x O (3) regime was provided by means of a free-air O(3) canopy exposure system. The hypotheses were tested that (1) gas exchange is negatively affected by O(3) and (2) the effects of O(3) are dose-dependent and thus the sizes of differences between treatments are positively related to COU. Gas exchange (light-saturated CO(2) uptake rate A(max), stomatal conductance g (s), maximum rate of carboxylation Vc (max), ribulose-1,5-bisphosphate turnover limited rate of photosynthesis J (max), CO(2) compensation point CP, apparent quantum yield of net CO(2) uptake AQ, carboxylation efficiency CE, day- and nighttime respiration) and chlorophyll fluorescence (electron transfer rate, ETR) were measured IN SITU on attached sun and shade leaves. Measurements were made periodically throughout the growing seasons of 2003 (an exceptionally dry year) and 2004 (a year with average rainfall). In 2004 Vc(max), J(max), and CE were lower in trees receiving 2 x O(3) compared with the ambient O(3) regime (1 x O(3)). Treatment differences in Vc (max), J (max), CE were rather small in 2004 (i.e., parameter levels were lower by 10 - 30 % in 2 x O(3) than 1 x O(3)) and not significant in 2003. In 2004 COU was positively correlated with the difference between treatments in A (max), g (s), and ETR (i.e., consistent with the dose-dependence of O(3)'s deleterious effects). However, in 2003, differences in A(max), g (s), and ETR between the two O(3) regimes were smaller at the end of the dry summer 2003 (i.e., when COU was greatest). The relationship of COU with effects on gas exchange can apparently be complex and, in fact, varied between years and within the growing season. In addition, high doses of O(3) did not always have significant effects on leaf gas exchange. In view of the key findings, both hypotheses were to be rejected.

Antioxidative defence of old growth beech (Fagus sylvatica) under double ambient O3 concentrations in a free-air exposure system.

In this study the influence of chronic free-air ozone exposure and of different meteorological conditions in the very dry year 2003 and the more humid year 2004 on the antioxidative system in sun and shade leaves of adult FAGUS SYLVATICA trees were investigated. Contents of ascorbate, glutathione, and alpha-tocopherol, as well as chloroplast pigments were determined under ambient (1 x O(3)) and double ambient (2 x O(3)) ozone concentrations. Ozone affected the antioxidative system in June and July, causing lower ascorbate contents in the apoplastic space, a more oxidized redox state of ascorbate and glutathione and an increase in pigment contents predominantly in the shade crown. For all measured parameters significant differences between the years were observed. In 2004 the redox state of ascorbate and glutathione was in a more reduced state and leaf contents of alpha-tocopherol, pigments of the xanthophyll cycle, beta-carotene, lutein, neoxanthin, and alpha-carotene were lower compared to 2003. Contents of total glutathione and chlorophyll a + b were increased in the second year. These results indicate a strong influence of the drought conditions in 2003 on the antioxidative system of beech overruling the ozone effects. Shade leaves showed lower contents of ascorbate in both years and the redox states of ascorbate and glutathione were more oxidized compared to sun leaves. Contents of photoprotective and accessory pigments generally were enhanced and the de-epoxidation state of the xanthophyll cycle was lower in the shade compared to the sun crown. Exhibiting less antioxidants shade leaves seem to be more sensitive against ozone than sun leaves.

Significance of ozone exposure for inter-annual differences in primary metabolites of old-growth beech (Fagus sylvatica L.) and Norway spruce (Picea abies L.) trees in a mixed forest stand.

The influence of long-term free-air ozone fumigation and canopy position on leaf contents of total glutathione, its redox state, non-structural proteins (NSP), soluble amino compounds, and total soluble sugars in old-growth beech (FAGUS SYLVATICA) and spruce (PICEA ABIES) trees were determined over a period of five years. Ozone fumigation had weak effects on the analysed metabolites of both tree species and significant changes in the contents of total glutathione, NSP, and soluble sugars were observed only selectively. Beech leaves were affected by crown position to a higher extent than spruce needles and exhibited lower contents of total glutathione and NSP and total soluble sugars, but enhanced contents of oxidised glutathione and amino compounds in the shade compared to the sun crown. Contents of total soluble sugars generally were decreased in shade compared to sun needles of spruce trees. Interspecific differences between beech and spruce were more distinct in the sun compared to the shade crown. Contents of total glutathione were increased whilst contents of amino compounds and total soluble sugars were lower in spruce needles compared to beech leaves. The metabolites determined showed individual patterns in the course of the five measurement years. Contents of total glutathione and its redox state correlated with air temperature and global radiation, indicating an important role for the antioxidant at low temperatures. Correlations of glutathione with instantaneous ozone concentrations seem to be a secondary effect. Differences in proteins and/or amino compounds in the inter-annual course are assumed to be a consequence of alterations in specific N uptake rates.

Tree internal signalling and defence reactions under ozone exposure in sun and shade leaves of European beech (Fagus sylvatica L.) trees.

The influence of free-air ozone (O(3)) fumigation on the levels of gene transcripts and compounds of defence and signalling were analysed in leaves of adult beech trees from the "Kranzberg Forest" research site in 2003 and 2004. This includes the precursor of the stress hormone ethylene, ACC (1-aminocyclopropane-1-carboxylic acid), conjugated salicylic acid, lignin content as well as of the expression level of genes connected with oxidative stress and stress signalling. At this site mature beech trees were exposed to an enhanced O(3) regime by a free-air O(3) canopy exposure system. Levels of conjugated ACC and conjugated salicylic acid in leaves were increased under O (3) fumigation whereas lignin content was only slightly enhanced. Quantitative real-time RT-PCR (qRT-PCR) was performed on transcripts of genes connected with lignin, salicylic acid, and ethylene formation, the shikimate pathway, abscisic acid biosynthesis as well as with the antioxidative system. Genes which showed O(3)-dependent increases included FSCOMT (caffeic-acid O-methyltransferase) connected with lignin formation, the stress response genes FSACS2 (ACC synthase) and FSPR1 (PR10 - pathogenesis-related protein), as well as FSNCED1 (9-cis-epoxicarotenoid dioxygenase), the rate-limiting enzyme of the ABA synthesis. For FSNCED1 expression level, a significant O(3) effect was found with an 8-fold (sun) and 7-fold (shade) induction in July 2003 and a 3-fold and 2.5-fold induction in July 2004. While the observed effects were not continuous, elevated O(3) is concluded to have the potential to disrupt the defence and signalling system.

Modelling ozone effects on adult beech trees through simulation of defence, damage, and repair costs: Implementation of the CASIROZ ozone model in the ANAFORE forest model.

Ozone affects adult trees significantly, but effects on stem growth are hard to prove and difficult to correlate with the primary sites of ozone damage at the leaf level. To simulate ozone effects in a mechanistic way, at a level relevant to forest stand growth, we developed a simple ozone damage and repair model (CASIROZ model) that can be implemented into mechanistic photosynthesis and growth models. The model needs to be parameterized with cuvette measurements on net photosynthesis and dark respiration. As the CASIROZ ozone sub-model calculates effects of the ozone flux, a reliable representation of stomatal conductance and therefore ozone uptake is necessary to allow implementation of the ozone sub-model. In this case study the ozone sub-model was used in the ANAFORE forest model to simulate gas exchange, growth, and allocation. A preliminary run for adult beech (FAGUS SYLVATICA) under different ozone regimes at the Kranzberg forest site (Germany) was performed. The results indicate that the model is able to represent the measured effects of ozone adequately, and to distinguish between immediate and cumulative ozone effects. The results further help to understand ozone effects by distinguishing defence from damage and repair. Finally, the model can be used to extrapolate from the short-term results of the field study to long-term effects on tree growth. The preliminary simulations for the Kranzberg beech site show that, although ozone effects on yearly growth are variable and therefore insignificant when measured in the field, they could become significant at longer timescales (above 5 years, 5 % reduction in growth). The model offers a possible explanation for the discrepancy between the significant effects on photosynthesis (10 to 30 % reductions simulated), and the minor effects on growth. This appears to be the result of the strong competition and slow growth of the Kranzberg forest, and the importance of stored carbon for the adult beech (by buffering effects on carbon gain). We finally conclude that inclusion of ozone effects into current forest growth and yield models can be an important improvement into their overall performance, especially when simulating younger and less dense forests.

Pacific Ocean and Japan Sea ecotypes of Japanese beech (Fagus crenata) differ in photosystem responses to continuous high light.

Two ecotypes of Japanese beech (Fagus crenata Blume), the Pacific Ocean type (PAO) and the Japan Sea type (JAS), show different responses to high solar irradiance. When PAO and JAS saplings were grown in continuous high-light (H), leaves of JAS became pale green. To elucidate this phenomenon, we investigated in vivo photochemistry based on pigment concentrations of Photosystem (PS) I and PS II and Western blot analysis. In JAS-H leaves, the amount of D1-protein decreased, resulting in decreases in the maximal quantum yield of PS II (F(v)/F(m)) and electron transport rate, whereas PAO-H leaves maintained high activities. The PS I photochemistry determined by measurement of P-700 photo-oxidation showed that the intersystem electron pool size was 1.4 times greater in JAS-H leaves than in PAO-H leaves. Furthermore, the re-reduction kinetics of P-700(+) showed that cyclic electron transport around PS I was 1.2 times faster in PAO-H leaves than in JAS-H leaves. Analysis of the area over the fluorescence induction kinetics indicated that the relative abundance of the PS IIalpha center increased in PAO-H leaves, whereas JAS leaves were observed to have low acclimation capacity to high light. These results demonstrate that PAO leaves possess acclimation mechanisms to continuous high light, whereas JAS leaves are more vulnerable to continuous high light, resulting in reduced leaf longevity owing to photoinhibition caused by increases in the intersystem electron pool size and suppression of photochemistry at the level of PS I and PS II.

Distribution of UV-shielding of the epidermis of sun and shade leaves of the beech (Fagus sylvatica L.) as monitored by multi-colour fluorescence imaging.

Plants can protect against damaging ultraviolet (UV) radiation by accumulating UV-absorbing substances in the epidermis of the leaves. Sun and shade leaves of a free standing beech tree (Fagus sylvatica L.) were studied for the differences in UV-shielding of the epidermis by means of multi-colour fluorescence images taken with UV and blue excitation. The distribution of the fluorescence intensity was detected over intact leaves in the emission maxima in the blue at 440 nm (F440), in the green at 520 nm (F520), in the red at 690 nm (F690) and in the far red at 740 nm (F740). Images of the logarithmic ratio between F690 excited in the blue and the UV (log ((B)F690/(UV)F690)) were calculated representing the relative absorption of UV in the epidermis and thus the degree of UV-shielding. It was found that UV-shielding is stronger for sun leaves than for shade leaves and better for the upper (adaxial) leaf side than for the lower (abaxial) leaf side of both leaf types. Within one leaf the highest value for the ratio log ((B)F690/(UV)F690) and thus the highest UV-shielding was found at the leaf rim which in broad leaves contains young tissue.

Investigating the European beech (Fagus sylvatica L.) leaf characteristics along the vertical canopy profile: leaf structure, photosynthetic capacity, light energy dissipation and photoprotection mechanisms.

Forest functionality and productivity are directly related to canopy light interception and can be affected by potential damage from high irradiance. However, the mechanisms by which leaves adapt to the variable light environments along the multilayer canopy profile are still poorly known. We explored the leaf morphophysiological and metabolic responses to the natural light gradient in a pure European beech (Fagus sylvatica L.) forest at three different canopy heights (top, middle and bottom). Structural adjustment through light-dependent modifications in leaf mass per area was the reason for most of the variations in photosynthetic capacity. The different leaf morphology along the canopy influenced nitrogen (N) partitioning, water- and photosynthetic N-use efficiency, chlorophyll (Chl) fluorescence and quali-quantitative contents of photosynthetic pigments. The Chl a to Chl b ratio and the pool of xanthophyll-cycle pigments (VAZ) increased at the highest irradiance, as well as lutein and ß-carotene. The total pool of ascorbate and phenols was higher in leaves of the top and middle canopy layers when compared with the bottom layer, where the ascorbate peroxidase was relatively more activated. The non-photochemical quenching was strongly and positively related to the VAZ/(Chl a + b) ratio, while Chl a/Chl b was related to the photochemical efficiency of photosystem II. Along the multilayer canopy profile, the high energy dissipation capacity of leaves was correlated to an elevated redox potential of antioxidants. The middle layer gave the most relevant contribution to leaf area index and carboxylation capacity of the canopy. In conclusion, a complex interplay among structural, physiological and biochemical traits drives the dynamic leaf acclimation to the natural gradients of variable light environments along the tree canopy profile. The relevant differences observed in leaf traits within the canopy positions of the beech forest should be considered for improving estimation of carbon fluxes in multilayer canopy models of temperate forests.

Photosynthetic traits of Siebold's beech seedlings in changing light conditions by removal of shading trees under elevated CO2.

The purpose of this study was to obtain basic information on acclimation capacity of photosynthesis in Siebold's beech seedlings to increasing light intensity under future elevated CO2 conditions. We monitored leaf photosynthetic traits of these seedlings in changing light conditions (before removal of shade trees, the year after removal of shade trees and after acclimation to open conditions) in a 10-year free air CO2 enrichment experiment in northern Japan. Elevated CO2 did not affect photosynthetic traits such as leaf mass per area, nitrogen content and biochemical photosynthetic capacity of chloroplasts (i.e. maximum rate of carboxylation and maximum rate of electron transport) before removal of the shade trees and after acclimation to open conditions; in fact, a higher net photosynthetic rate was maintained under elevated CO2 . However, in the year after removal of the shade trees, there was no increase in photosynthesis rate under elevated CO2 conditions. This was not due to photoinhibition. In ambient CO2 conditions, leaf mass per area and nitrogen content were higher in the year after removal of shade trees than before, whereas there was no increase under elevated CO2 conditions. These results indicate that elevated CO2 delays the acclimation of photosynthetic traits of Siebold's beech seedlings to increasing light intensity.