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As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (gs) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (udaily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, udaily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and gs recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and gs. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of udaily at the crown base compared with DBH level upon watering. However, udaily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g., udaily, ΨPD and gs, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored.

Phloem sustains maintenance and growth processes through transport of sugars from source to sink organs. Under low water availability, tree functioning is impaired, i.e., growth/photosynthesis decline and phloem transport may be hindered. In a 3-year throughfall exclusion (TE) experiment on mature European beech (Fagus sylvatica L.) we conducted 13CO2 branch labeling to investigate translocation of recently fixed photoassimilates under experimental drought over 2 years (2015 and 2016). We hypothesized (H1) that mean residence time of photoassimilates in leaves (MRT) increases, whereas (H2) phloem transport velocity (Vphloem) decreases under drought. Transport of carbohydrates in the phloem was assessed via δ13C of CO2 efflux measured at two branch positions following 13CO2 labeling. Pre-dawn water potential (ΨPD) and time-integrated soil water deficit (iSWD) were used to quantify drought stress. The MRT increased by 46% from 32.1 ± 5.4 h in control (CO) to 46.9 ± 12.3 h in TE trees, supporting H1, and positively correlated (P < 0.001) with iSWD. Confirming H2, Vphloem in 2016 decreased by 47% from 20.7 ± 5.8 cm h-1 in CO to 11.0 ± 2.9 cm h-1 in TE trees and positively correlated with ΨPD (P = 0.001). We suggest that the positive correlation between MRT and iSWD is a result of the accumulation of osmolytes maintaining cell turgor in the leaves under longer drought periods. Furthermore, we propose that the positive correlation between Vphloem and ΨPD is due to a lower water uptake of phloem conduits from surrounding tissues under increasing drought leading to a higher phloem sap viscosity and lower Vphloem. The two mechanisms increasing MRT and reducing Vphloem respond differently to low water availability and impair trees' carbon translocation under drought.

Under drought, potential plant death from depletion of carbon (C) stores, referred to as carbon starvation, is thought to result from negative carbon balance during (partial) stomatal closure. As evidence for C starvation is rare for mature trees, we investigated the C dynamics in mature beech and spruce under drought, focusing on non-structural carbohydrates (NSC) as an indicator of their C balance. Trees were exposed to complete exclusion of precipitation throughfall over two consecutive growing seasons. We assessed NSC concentrations during the early dormant season in leaves, twigs, stem phloem/xylem, coarse and fine roots. Up-scaling to whole-tree NSC pools was achieved using a process-based single-tree model (“BALANCE”), estimating tree biomass increment. While there were distinct differences in NSC concentration among different tissues in both beech and spruce (root

Hydraulic redistribution (HR) of soil water through plant roots is a crucial phenomenon improving the water balance of plants and ecosystems. It is mostly described under severe drought, and not yet studied under moderate drought. We tested the potential of HR under moderate drought, hypothesizing that (H1) tree species redistribute soil water in their roots even under moderate drought and that (H2) neighboring plants are supported with water provided by redistributing plants. Trees were planted in split-root systems with one individual (i.e., split-root plant, SRP) having its roots divided between two pots with one additional tree each. Species were 2- to 4-year-old English oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). A gradient in soil water potential (ψsoil) was established between the two pots (-0.55 ± 0.02 MPa and -0.29 ± 0.03 MPa), and HR was observed by labeling with deuterium-enriched water. Irrespective of species identity, 93% of the SRPs redistributed deuterium enriched water from the moist to the drier side, supporting H1. Eighty-eight percent of the plants in the drier pots were deuterium enriched in their roots, with 61 ± 6% of the root water originating from SRP roots. Differences in HR among species were related to their root anatomy with diffuse-porous xylem structure in both beech and-opposing the stem structure-oak roots. In spruce, we found exclusively tracheids. We conclude that water can be redistributed within roots of different tree species along a moderate ψsoil gradient, accentuating HR as an important water source for drought-stressed plants, with potential implications for ecohydrological and plant physiological sciences. It remains to be shown to what extent HR occurs under field conditions in Central Europe.

Vegetation responds to drought through a complex interplay of plant hydraulic mechanisms, posing challenges for model development and parameterization. We present a mathematical model that describes the dynamics of leaf water-potential over time while considering different strategies by which plant species regulate their water-potentials. The model has two parameters: the parameter λ describing the adjustment of the leaf water potential to changes in soil water potential, and the parameter Δψww describing the typical ‘well-watered’ leaf water potentials at non-stressed (near-zero) levels of soil water potential. Our model was tested and calibrated on 110 time-series datasets containing the leaf- and soil water potentials of 66 species under drought and non-drought conditions. Our model successfully reproduces the measured leaf water potentials over time based on three different regulation strategies under drought. We found that three parameter sets derived from the measurement data reproduced the dynamics of 53% of an drought dataset, and 52% of a control dataset [root mean square error (RMSE) < 0.5 MPa)]. We conclude that, instead of quantifying water-potential-regulation of different plant species by complex modeling approaches, a small set of parameters may be sufficient to describe the water potential regulation behavior for large-scale modeling. Thus, our approach paves the way for a parsimonious representation of the full spectrum of plant hydraulic responses to drought in dynamic vegetation models.

Key message Under severe drought, growth of Norway spruce suffered much more than European beech. Norway spruce benefited from growing in the environment of beech, and both species acclimated slightly to 5 years of experimentally extended drought. Abstract Recent studies show that the detrimental effects of drought on stand growth are mitigated when the stand contains mixed tree species. We analysed the growth responses of Norway spruce and European beech to episodic and experimentally extended drought in intra- and inter-specific mature stands. We used annual diameter growth records dating back to 1998 to determine the impact of the natural episodic drought in 2003 and 2015. To analyse extended drought, spruce and beech trees were exposed to extreme drought under automatic throughfall exclusion roofs from 2014 to 2018. The growth of spruce in an inter-specific environment with beech was 20–50% less affected by natural episodic drought compared with an intra-specific constellation. When beech grew in an inter-specific environment, it was by 23% more affected by drought compared to intra-specific conditions, but seemed to recover faster. The induced drought from 2014 to 2018 resulted in a strong growth reduction in the first year particularly for spruce, followed by a slight acclimation to the dry conditions. Beech acclimated and recovered faster than spruce across all growing conditions, while spruce only acclimatized faster in the environment of beech. Both species showed a higher mortality under induced drought compared with the controls; for spruce, the mortality rate was fivefold higher compared to the long-term mortality. The long-term moderate-growth stabilization and the growth increase after the 5-year exposure to drought suggest a gradual acclimation to drought by beech. The resistance and acclimation to drought of spruce when growing in mixture should be considered when designing resource efficient and productive mixed conifer-broadleaved stands for future climates.

Hydraulic redistribution (HR) can buffer drought events of tree individuals, however, its relevance for neighbouring trees remains unclear. Here, we quantified HR to neighbouring trees in single- and mixed-species combinations. We hypothesized that uptake of HR water positively correlates with root length, number of root tips and root xylem hydraulic conductivity and that neighbours in single-species combinations receive more HR water than in phylogenetic distant mixed-species combinations. In a split-root experiment, a sapling with its roots split between two pots redistributed deuterium labelled water from a moist to a dry pot with an additional tree each. We quantified HR water received by the sapling in the dry pot for six temperate tree species. After 7 days, one quarter of the water in roots (2.1 ± 0.4 ml), stems (0.8 ± 0.2 ml) and transpiration (1.0 ± 0.3 ml) of the drought stressed sapling originated from HR. The amount of HR water transpired by the receiving plant stayed constant throughout the experiment. While the uptake of HR water increased with root length, species identity did not affect HR as saplings of Picea abies ((L.) Karst) and Fagus sylvatica (L.) in single- and mixed-species combinations received the same amount of HR water.

The (13/12) C ratio in plant roots is likely dynamic depending on root function (storage versus uptake), but to date, little is known about the effect of season and root order (an indicator of root function) on the isotopic composition of C-rich fractions in roots. To address this, we monitored the stable isotopic composition of one evergreen (Picea abies) and one deciduous (Fagus sylvatica), tree species' roots by measuring δ(13) C of bulk, respired and labile C, and starch from first/second and third/fourth order roots during spring and fall root production periods. In both species, root order differences in δ(13) C were observed in bulk organic matter, labile, and respired C fractions. Beech exhibited distinct seasonal trends in δ(13) C of respired C, while spruce did not. In fall, first/second order beech roots were significantly depleted in (13) C, whereas spruce roots were enriched compared to higher order roots. Species variation in δ (13) C of respired C may be partially explained by seasonal shifts from enriched to depleted C substrates in deciduous beech roots. Regardless of species identity, differences in stable C isotopic composition of at least two root order groupings (first/second, third/fourth) were apparent, and should hereafter be separated in belowground C-supply-chain inquiry.

As long-lived organisms, trees use resources to support both growth and below- and aboveground trophic interactions. Resources fluctuate in relation to periods of growth cease that are regulated by internal and external factors, and these fluctuations feed backs to trophic partners. Some major forest trees display an endogenous growth rhythm, and related pulses of variation in allocation of resources have been detected. As this trait makes it possible to separate growth into defined phases, it offers an opportunity to disentangle the intermingled complex regulation of growth and multitrophic interactions in trees. We present “TrophinOak”, a platform using microcuttings of pedunculated oak, a tree that displays endogenous rhythmic growth characterized by alternating shoot and root growth flushes. We select seven beneficial or detrimental above- and belowground partners including animals (Lymantria dispar, Pratylenchus penetrans, Protaphorura armata), fungi (Piloderma croceum, Microsphaera alphitoides, Phytophthora quercina) and bacteria (Streptomyces sp.), to synthesize bi- and tripartite trophic interactions, including ectomycorrhizal symbioses, and monitor fluctuations of carbon and nitrogen allocation as well as plant gene expression at distinct phases of oak growth. We use this model to identify and resolve the experimental challenges inherent in synthesizing diverse types of associations in a common microcosm system, in labeling plants with stable N and C isotopes and in analyzing transcripts in a non-model plant, a process which requires generating a specific contig library. We develop hypotheses and experimental design to test them in order to identify core mechanisms that help trees to modulate their own development and their multitrophic interactions for optimizing their long term performance in their environment. First results constitute a proof of concept that the platform works and enables us to test the hypotheses.

Absorptive fine roots are an important driver of soil biogeochemical cycles. Yet, the spatio-temporal dynamics of those roots in the presence of neighboring species remain poorly understood. The aim of this study was to analyze shifts in absorptive fine-root traits in monoculture or mixtures of Fagus sylvatica [L.] and Picea abies [L.] Karst. We hypothesized that root competition would be higher under single-species than mixed-species interactions, leading to changes in (i) root survivorship, diameter and respiration and (ii) spatio-temporal patterns of root growth and death. Using minirhizotron methods, we monitored the timing and location of absorptive fine-root growth and death at an experimental forest in southern Germany from 2011 to 2013. We also measured root respiration in the spring and fall seasons of 2012 and 2013. Our findings show that the absorptive fine roots of F. sylvatica had a 50% higher risk of root mortality and higher respiration rates in the single-species compared to mixed-species zones. These results support our hypothesis that root competition is less intense for F. sylvatica in mixture versus monoculture. We were unable to find confirmation for the same hypothesis for P. abies. To analyze spatio-temporal patterns of absorptive fine-root production and mortality, we used a mixed-effects model considering root depth (space) and seasons (time) simultaneously. This analysis showed that F. sylvatica shifts root production towards shallower soil layers in mixed-species stands, besides significant seasonal fluctuations in root production depths for both species. Ultimately, the impact of neighbor species identity on root traits observed in this study has important implications for where, when and how fast root-facilitated carbon cycling takes place in single-species versus mixed-species forests. In addition, our study highlights the need for inclusion of absorptive fine-root spatio-temporal dynamics when examining belowground plant interactions and biogeochemical cycles.

International audience; nitori Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die-off events have severe consequences for ecosystem services, biophysical and biogeochemical land-atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global mong network of forest mortality. In light of recent and anticipated large forest die-off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought-induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level.

Highlight Increased resource availability has no impact on endogenous rhythmic growth and related allocation shifts in oak trees; an internal clock regulates core genes during shoot and root growth cessation phases., Common oak trees display endogenous rhythmic growth with alternating shoot and root flushes. To explore the mechanisms involved, microcuttings of the Quercus robur L. clone DF159 were used for 13C/15N labelling in combination with RNA sequencing (RNASeq) transcript profiling of shoots and roots. The effect of plant internal resource availability on the rhythmic growth of the cuttings was tested through inoculation with the ectomycorrhizal fungus Piloderma croceum. Shoot and root flushes were related to parallel shifts in above- and below-ground C and, to a lesser extent, N allocation. Increased plant internal resource availability by P. croceum inoculation with enhanced plant growth affected neither the rhythmic growth nor the associated resource allocation patterns. Two shifts in transcript abundance were identified during root and shoot growth cessation, and most concerned genes were down-regulated. Inoculation with P. croceum suppressed these transcript shifts in roots, but not in shoots. To identify core processes governing the rhythmic growth, functions [Gene Ontology (GO) terms] of the genes differentially expressed during the growth cessation in both leaves and roots of non-inoculated plants and leaves of P. croceum-inoculated plants were examined. Besides genes related to resource acquisition and cell development, which might reflect rather than trigger rhythmic growth, genes involved in signalling and/or regulated by the circadian clock were identified. The results indicate that rhythmic growth involves dramatic oscillations in plant metabolism and gene regulation between below- and above-ground parts. Ectomycorrhizal symbiosis may play a previously unsuspected role in smoothing these oscillations without modifying the rhythmic growth pattern.

Summary The root-rot pathogen Phytophthora quercina is a key determinant of oak decline in Europe. The susceptibility of pedunculate oak (Quercus robur) to this pathogen has been hypothesized to depend on the carbon availability in roots as an essential resource for defense. Microcuttings of Q. robur undergo an alternating rhythm of root and shoot growth. Inoculation of mycorrhizal (Piloderma croceum) and nonmycorrhizal oak roots with P. quercina was performed during both growth phases, that is, root flush (RF) and shoot flush (SF). Photosynthetic and morphological responses as well as concentrations of nonstructural carbohydrates (NSC) were analyzed. Infection success was quantified by the presence of pathogen DNA in roots. Concentrations of NSC in roots depended on the alternating root/shoot growth rhythm, being high and low during RF and SF, respectively. Infection success was high during RF and low during SF, resulting in a significantly positive correlation between pathogen DNA and NSC concentration in roots, contrary to the hypothesis. The alternating growth of roots and shoots plays a crucial role for the susceptibility of lateral roots to the pathogen. NSC availability in oak roots has to be considered as a benchmark for susceptibility rather than resistance against P. quercina.

We review causes of synergies in mixed-species stands, develop guiding hypotheses for revealing their mechanisms and present a rainfall exclusion experiment along with a transect (KROOF) for exploring drought effects. While monocultures have dominated forest research and practice in the past, in face of growing resource scarcity and climate change, mixed-species stands are on the advance. Long-term observations show that mixed-species stands frequently over-yield monocultures, and they further suggest that the over-yielding is often higher on poor than on fertile sites and in low-growth than in high-growth years. However, the underlying causes have not yet been clarified. We start with a review of, among others, hydraulic redistribution, complementary eco-physiological traits, and ectomycorrhizal networks as possible causes behind the observed productivity gains in mixed-species stands. Then, we develop guiding hypotheses for further exploration of synergies in mixed-species stands. Finally, we introduce into the concept and model approach of the rainfall exclusion experiment for examining the role of water supply in mixed versus pure forest stands of spruce and beech. At the Kranzberg site, six plots are equipped with water retention roofs, which only close during rain events. The remaining six plots serve as non-roofed control. Together with the rainfall exclusion experiment, an ecological gradient with five sites extending through SE-Germany forms the “Kranzberg Roof Experiment” (in short KROOF). Kranzberg Forest is a part of this gradient from moist to dry conditions, with each site providing mixed and pure associations each of spruce and beech. The ecophysiological approach will be complemented by tree ring analysis and modelling of productivity of the tree associations under intense drought.

Our study provides evidence that neither elevated CO 2 nor elevated O 3 alters the positive asymmetric competition for light and the symmetric competition for water among beech and spruce individuals grown in monoculture. We conclude that the mechanism of competition (i.e. symmetric/asymmetric) above (e.g shading or overtopping effect) and belowground (e.g. non-preemption or foraging) rather than abiotic treatments such as elevated CO 2 , O 3 and CO 2 /O 3 regimes, plays a dominant role for ensuring competitive success among tree saplings. Despite numerous studies conducted on plant responses to increasing CO2 and O3 concentrations, there is still a gap in understanding on how these gasses would affect the mode of competition (e.g., the ability by which larger and smaller plants capture resources) at the individual level of intra-specific beech and spruce saplings. Using empirical data and simulations from the plant-growth model PLATHO, we analyzed underlying mechanisms of competition and extrapolated effects beyond the time span of the experiment. We hypothesized that among juvenile beech and spruce trees planted in monoculture, +CO2 would diminish the positive asymmetric competition for light. Conversely, +O3 would enhance this outcome. In addition, we hypothesized that the symmetric mode of competition belowground for water would remain unchanged, irrespective of +CO2 and/or +O3 treatments. Our results showed that +CO2 and/or +O3 treatments did not alter the mode of competition aboveground for light. Conversely, we accepted our hypothesis that the mode of competition for water would remain unchanged under both treatments. Overall, we conclude that neither +CO2 nor +O3 alters the positive asymmetric competition for light and the symmetric competition for water among beech and spruce individuals grown in monoculture. We further conclude that competitive mechanism above (e.g., shading or overtopping effect) and belowground (e.g., non-preemption or foraging) rather than abiotic treatments, such as elevated CO2, O3 and CO2/O3 regimes, plays a dominant role for ensuring competitive success among tree saplings.

Leaf photosynthesis of the sensitive plant Mimosa pudica displays a transient knockout in response to electrical signals induced by heat stimulation. This study aims at clarifying the underlying mechanisms, in particular, the involvement of respiration. To this end, leaf gas exchange and light reactions of photosynthesis were assessed under atmospheric conditions largely eliminating photorespiration by either elevated atmospheric CO2 or lowered O2 concentration (i.e. 2000 μmol mol(-1) or 1%, respectively). In addition, leaf gas exchange was studied in the absence of light. Under darkness, heat stimulation caused a transient increase of respiratory CO2 release simultaneously with stomatal opening, hence reflecting direct involvement of respiratory stimulation in the drop of the net CO2 uptake rate. However, persistence of the transient decline in net CO2 uptake rate under illumination and elevated CO2 or 1% O2 makes it unlikely that photorespiration is the metabolic origin of the respiratory CO2 release. In conclusion, the transient knockout of net CO2 uptake is at least partially attributed to an increased CO2 release through mitochondrial respiration as stimulated by electrical signals. Putative CO2 limitation of Rubisco due to decreased activity of carbonic anhydrase was ruled out as the photosynthesis effect was not prevented by elevated CO2 .

Summary Oaks (Quercus spp.), which are major forest trees in the northern hemisphere, host many biotic interactions, but molecular investigation of these interactions is limited by fragmentary genome data. To date, only 75 oak expressed sequence tags (ESTs) have been characterized in ectomycorrhizal (EM) symbioses. We synthesized seven beneficial and detrimental biotic interactions between microorganisms and animals and a clone (DF159) of Quercus robur. Sixteen 454 and eight Illumina cDNA libraries from leaves and roots were prepared and merged to establish a reference for RNASeq transcriptomic analysis of oak EMs with Piloderma croceum. Using the Mimicking Intelligent Read Assembly (MIRA) and Trinity assembler, the OakContigDF159.1 hybrid assembly, containing 65 712 contigs with a mean length of 1003 bp, was constructed, giving broad coverage of metabolic pathways. This allowed us to identify 3018 oak contigs that were differentially expressed in EMs, with genes encoding proline-rich cell wall proteins and ethylene signalling-related transcription factors showing up-regulation while auxin and defence-related genes were down-regulated. In addition to the first report of remorin expression in EMs, the extensive coverage provided by the study permitted detection of differential regulation within large gene families (nitrogen, phosphorus and sugar transporters, aquaporins). This might indicate specific mechanisms of genome regulation in oak EMs compared with other trees.

Climate simulations anticipate an increase in mean summer temperature with synchronous decrease in summer precipitation during the course of the current century in Central Europe. As a consequence, transpiration of forest trees and stands may be altered along with soil water availability. In this study, the effect of reduced plant water availability to conifers was investigated into an open Pinus sylvestris forest (Erico-Pinetum typicum; P. sylvestris 60 %, Picea abies 20 %; and Larix decidua 20 %) within the inner alpine dry valley of the Inn River in Tyrol, Austria. For reducing plant water availability, we installed a transparent roof construction above the forest floor to prevent precipitation to reach the soil. The roofed area covered 240 m² and included 10 trees. A respective number of 11 trees served as controls in the absence of any manipulation. Roofing significantly reduced plant water availability as indicated in lower predawn needle water potentials. Sap flow density (Q s) was 63, 47, and 24 % lower in roofed P. sylvestris, P. abies, and L. decidua trees, respectively, as compared to control trees. Our findings suggest that P. sylvestris and P. abies behaves “isohydric” as they close their stomata relatively early under conditions of reduced plant water availability and thus stabilize their water relations, whereas L. decidua behaves “anisohydric” and maintains high transpiration rates.

• The CO(2) efflux of adult trees is supplied by recent photosynthates and carbon (C) stores. The extent to which these C pools contribute to growth and maintenance respiration (R(G) and R(M), respectively) remains obscure. • Recent photosynthates of adult beech (Fagus sylvatica) and spruce (Picea abies) trees were labeled by exposing whole-tree canopies to (13) C-depleted CO(2). Label was applied three times during the year (in spring, early summer and late summer) and changes in the stable C isotope composition (δ(13) C) of trunk and coarse-root CO(2) efflux were quantified. • Seasonal patterns in C translocation rate (CTR) and fractional contribution of label to CO(2) efflux (F(Label-Max)) were found. CTR was fastest during early summer. In beech, F(Label-Max) was lowest in spring and peaked in trunks during late summer (0.6 ± 0.1, mean ± SE), whereas no trend was observed in coarse roots. No seasonal dynamics in F(Label-Max) were found in spruce. • During spring, the R(G) of beech trunks was largely supplied by C stores. Recent photosynthates supplied growth in early summer and refilled C stores in late summer. In spruce, CO(2) efflux was constantly supplied by a mixture of stored (c. 75%) and recent (c. 25%) C. The hypothesis that R(G) is exclusively supplied by recent photosynthates was rejected for both species.

Respiration is a substantial driver of carbon (C) flux in forest ecosystems and stable C isotopes provide an excellent tool for its investigation. We studied seasonal dynamics in δ¹³C of CO₂ efflux (δ¹³C(E)) from non-leafy branches, upper and lower trunks and coarse roots of adult trees, comparing deciduous Fagus sylvatica (European beech) with evergreen Picea abies (Norway spruce). In both species, we observed strong and similar seasonal dynamics in the δ¹³C(E) of above-ground plant components, whereas δ¹³C(E) of coarse roots was rather stable. During summer, δ¹³C(E) of trunks was about -28.2‰ (Beech) and -26.8‰ (Spruce). During winter dormancy, δ¹³C(E) increased by 5.6-9.1‰. The observed dynamics are likely related to a switch from growth to starch accumulation during fall and remobilization of starch, low TCA cycle activity and accumulation of malate by PEPc during winter. The seasonal δ¹³C(E) pattern of branches of Beech and upper trunks of Spruce was less variable, probably because these organs were additionally supplied by winter photosynthesis. In view of our results and pervious studies, we conclude that the pronounced increases in δ¹³C(E) of trunks during the winter results from interrupted access to recent photosynthates.

Stable carbon (C) isotopes, in particular employed in labeling experiments, are an ideal tool to broaden our understanding of C dynamics in trees and forest ecosystems. Here, we present a free-air exposure system, named isoFACE, designed for long-term stable C isotope labeling in the canopy of 25 m tall forest trees. Labeling of canopy air was achieved by continuous release of CO2 with a δ13C of −46.9‰. To this end, micro-porous tubes were suspended at c. 1 m distance vertically through the canopy, minimizing CO2 gradients from the exterior to the interior and allowing for C labeling exposure during periods of low wind speed. Target for CO2 concentration ([CO2]) increase was ambient +100 μmol mol−1. Canopy [CO2] stayed within 10% of the target during more than 57% of the time and resulted in a drop of δ13C in canopy air by 7.8‰. After 19 labeling days about 50% of C in phloem sugars and stem CO2 efflux were turned over and 20–30% in coarse root CO2 efflux and soil CO2. The isoFACE system successfully altered δ13C of canopy air for studying turn-over of C pools in forest trees and soils, highlighting their slow turn-over rates.

In a lysimeter study with young beech trees, the effects of elevated ozone concentration on the decomposition and fate of nitrogen in 15N‐labeled leaf litter were analyzed after one growing season. Nitrogen in the litter was dominated by a relatively inert, residual fraction, but easily decomposable nitrogen was present in substantial amounts. Nitrogen loss was significantly higher at twice‐ambient ozone which was largely attributed to an enhanced mobilization of residual nitrogen. Enhanced mobilization of nitrogen from litter at twice‐ambient ozone exposure resulted in additional 15N incorporation into the soil down to 30 cm depth. Only 0.41–0.62% of the nitrogen in the litter was incorporated into plant material at both ozone concentrations. Twice‐ambient ozone exposure changed the distribution of the nitrogen taken up from litter inside the beech trees in favor of the shoot, where it may have been used in biosynthetic processes required for defense reactions.

Ecuadorian mountain rainforests are declining dramatically due to deforestation. Exploitation of remaining forests has led to low abundances of native, valuable timber species. Enrichment planting of selected native tree species into forest gaps is a strategy that may increase their abundance and maintain biodiversity. However, the development of successful planting strategies requires knowledge of environmental demands on, and ecological requirements of, native species during their establishment. This knowledge is currently lacking for mid- and late-successional species in Central American forests. Two deciduous, mid-successional (Cedrela montana, Tabebuia chrysantha) and two evergreen, late-successional native tree species (Nectandra membranacea, Podocarpus sprucei) were planted into felling gaps. Photosynthetic performance and growth in height of these species were assessed along light gradients during seedling establishment to test whether species-specific light responses were related to plant successional traits. Both mid-successional species benefited from higher light levels in gaps up to 30% canopy openness60°. In larger gaps, C. montana exhibited a significant decline in growth. As expected, growth of the late-successional species was only marginally increased at higher light levels. Nevertheless, the photosynthetic apparatus of N. membranacea displayed rapid acclimation to higher light conditions in gaps. Plant response to felling gaps may not always be predicted based on successional status. Our results suggest that the four investigated species may coexist in the same gap by occupying different niches along light gradients. This arrangement may offer an ecological basis to increase the abundance of valuable timber species through enrichment planting in Ecuador mountain rainforests.

The impact of chronic free air ozone (O3) exposure and belowground pathogen stress on growth and total biomass development of young beech trees (Fagus sylvatica L.) was investigated in a lysimeter study. Plants were growing during four years under ambient or elevated atmospheric O3 concentrations. Additionally, in the last vegetation period the root rot pathogen Phytophthora citricola was introduced to study the interaction of ozone exposure and pathogen stress in the soil-plant system. A complete harvest at the end of the experiment enabled for the first time the assessment of fine and coarse root biomass of individual trees with a high vertical resolution down to two meter depth. Plant growth was significantly reduced by elevated ozone but not affected by P. citricola. Biomass partitioning between fine and coarse roots as well as vertical root distribution were significantly affected by both factors, whereas changes in root/shoot biomass ratio were not observed.

Combining measurements of electric potential and pH with such of chlorophyll fluorescence and leaf gas exchange showed heat stimulation to evoke an electrical signal (propagation speed: 3-5 mm s(-1)) that travelled through the leaf while reducing the net CO(2) uptake rate and the photochemical quantum yield of both photosystems (PS). Two-dimensional imaging analysis of the chlorophyll fluorescence signal of PS II revealed that the yield reduction spread basipetally via the veins through the leaf at a speed of 1.6 +/- 0.3 mm s(-1) while the propagation speed in the intervein region was c. 50 times slower. Propagation of the signal through the veins was confirmed because PS I, which is present in the bundle sheath cells around the leaf vessels, was affected first. Hence, spreading of the signal along the veins represents a path with higher travelling speed than within the intervein region of the leaf lamina. Upon the electrical signal, cytoplasmic pH decreased transiently from 7.0 to 6.4, while apoplastic pH increased transiently from 4.5 to 5.2. Moreover, photochemical quantum yield of isolated chloroplasts was strongly affected by pH changes in the surrounding medium, indicating a putative direct influence of electrical signalling via changes of cytosolic pH on leaf photosynthesis.

Based on the growth-differentiation balance theory (GDB) and the influence of tropospheric ozone (O3) on plants, we hypothesized that pre-conditioning with elevated O3 reduces adverse effects of the root rot pathogen Phytophthora citricola Sawada. To this end a 2-year phytotron study with juvenile European beech (Fagus sylvatica L.) and (Picea abies [L.] Karst.) grown in mixture was performed. The hypothesis was tested on phenological, leaf and root morphological as well as physiological aspects of plant performance. Contrasting with spruce, elevated O3 limited leaf and root biomass development, photosynthetic performance and N uptake of beech. The growth limitation by O3 conveyed increased resistance in beech against the pathogen. Conversely, spruce displayed enhanced susceptibility in the combined O3/P. citricola treatment. The hypothesis was supported in the case of beech rather than spruce. Nevertheless, conclusions support GDB regarding the trade-off between growth and stress defense, although compliance appears to be species-specific.

Saplings of Fagus sylvatica and Picea abies were grown under conditions of intra and interspecific competition in a 2-year phytotron study under combinations of ambient and elevated ozone (+O3 which is 2 × O3, but

BACKGROUND AND AIMS Patterns and variations in concentration of carbon-based secondary compounds in plant tissues have been explained by means of different complementary and, in some cases, contradictory plant defence hypotheses for more than 20 years. These hypotheses are conceptual models which consider environmental impacts on plant internal demands. In the present study, a mathematical model is presented, which converts and integrates the concepts of the 'Growth-Differentiation Balance' hypothesis and the 'Protein Competition' model into a dynamic plant growth model, that was tested with concentration data of polyphenols in leaves of juvenile apple, beech and spruce trees. The modelling approach is part of the plant growth model PLATHO that considers simultaneously different environmental impacts on the most important physiological processes of plants. METHODS The modelling approach for plant internal resource allocation is based on a priority scheme assuming that growth processes have priority over allocation to secondary compounds and that growth-related metabolism is more strongly affected by nitrogen deficiency than defence-related secondary metabolism. KEY RESULTS It is shown that the model can reproduce the effect of nitrogen fertilization on allocation patterns in apple trees and the effects of elevated CO(2) and competition in juvenile beech and spruce trees. The analysis of model behaviour reveals that large fluctuations in plant internal availability of carbon and nitrogen are possible within a single vegetation period. Furthermore, the model displays a non-linear allocation behaviour to carbon-based secondary compounds. CONCLUSIONS The simulation results corroborate the underlying assumptions of the presented modelling approach for resource partitioning between growth-related primary metabolism and defence-related secondary metabolism. Thus, the dynamical modelling approach, which considers variable source and sink strengths of plant internal resources within different phenological growth stages, presents a successful translation of existing concepts into a dynamic mathematical model.

In Mimosa pudica L., heat stimulation triggers leaflet folding in local, neighbouring and distant leaves. Stomatal movements were observed microscopically during this folding reaction and electrical potentials, chlorophyll fluorescence, and leaf CO(2)/H(2)O-gas exchange were measured simultaneously. Upon heat stimulation of a neighbouring pinna, epidermal cells depolarized and the stomata began a rapid and pronounced transient opening response, leading to an approximately 2-fold increase of stomatal aperture within 60 s. At the same time, net CO(2) exchange showed a pronounced transient decrease, which was followed by a similar drop in photochemical quantum yield at photosystem (PS) II. Subsequently, CO(2)-gas exchange and photochemical quantum yield recovered and stomata closed partly or completely. The transient and fast stomatal opening response is interpreted as a hydropassive stomatal movement caused by a sudden loss of epidermal turgor. Thus, epidermal cells appear to respond in a similar manner to heat-induced signals as the pulvinar extensor cells. The subsequent closing of the stomata confirms earlier reports that stomatal movements can be induced by electrical signals. The substantial delay (several minutes) of guard cell turgor loss compared with the immediate response of the extensor and epidermal cells suggests a different, less direct mechanism for transmission of the propagating signal to the guard cells.

To gain an understanding of the role of electrical signaling in trees, poplar (Populus trichocarpa, Populus tremula × P. tremuloides) shoots were stimulated by chilling as well as flaming. Two kinds of signal propagation were detected by microelectrode measurements (aphid technique) in the phloem of leaf veins: (1) basipetal, short-distance signaling that led to rapid membrane hyperpolarization caused by K+-efflux within the leaf lamina; and (2) acropetal, long-distance signaling that triggered depolarization of the membrane potential in the leaf phloem. In the latter, the depolarizing signals travel across the stem from the manipulated leaves to adjacent leaves where the net CO2 uptake rate is temporarily depressed toward compensation. With regard to photosystem II, both heat-induced long-distance and short-distance signaling were investigated using two-dimensional “imaging” analysis of chlorophyll fluorescence. Both types of signaling significantly reduced the quantum yield of electron transport through photosystem II. Imaging analysis revealed that the signal that causes yield reduction spreads through the leaf lamina. Coldblocking of the stem proved that the electrical signal transmission via the phloem becomes disrupted, causing the leaf gas exchange to remain unaffected. Calcium-deficient trees showed a marked contrast inasmuch as the amplitude of the electrical signal was distinctly reduced, concomitant with the absence of a significant response in leaf gas exchange upon flame wounding. In summary, the above results led us to conclude that calcium as well as potassium is involved in the propagation of phloem-transmitted electrical signals that evoke specific responses in the photosynthesis of leaves.

To test the involvement of ethylene in mediating ozone- induced cell death and leaf loss in European beech ( Fagus sylvatica L.), tree seedlings were exposed to proportionally increased or decreased field ozone levels for up to 6 months. Ozone treatment caused cell death and acceler- ated leaf loss at higher than ambient levels, but had only minor effects at ambient and no effects at subambient ozone levels. The emission of ethylene, the levels of its precursor, 1-aminocyclopropane-1-carboxylate (ACC), and mRNA levels of specific ACC synthase ( FS-ACS2 ) and ACC oxidase ( FS-ACO1 ) isoforms showed a persistent increase and preceded cell death by approximately 2 weeks. Inhibition of ethylene biosynthesis led to reduced lesion formation whereas application of ACC accelerated ozone- induced cell death and leaf loss. Similar results were obtained when adult beech trees were exposed to 2 ¥ ¥ ¥ ozone by a whole tree free-air canopy exposure system. The results suggest a role of ethylene in amplifying ozone effects under field conditions in this major European broad-leaved tree species.

Summary •A 2-yr phytotron study was conducted to investigate the intra- and inter-specific competitive behaviour of juvenile beech ( Fagus sylvatica ) and spruce ( Picea abies ). Competitiveness was analysed by quantifying the resource budgets that occur along structures and within occupied space of relevance for competitive interaction. • Ambient and elevated CO 2 and ozone (O 3 ) regimes were applied throughout two growing seasons as stressors for provoking changes in resource budgets, growth and allocation to facilitate the competition analysis. The hypothesis tested was that the ability to sequester space at low structural cost will determine the competitive success. • Spruce was a stronger competitor than beech, as displayed by its higher aboveground biomass increments in mixed culture compared with monoculture. A crucial factor in the competitive success of spruce was its ability to enlarge crown volume at low structural costs, supporting the hypothesis. • Interspecific competition with spruce resulted in a size-independent readjustment of above-ground allocation in beech (reduced leaf : shoot biomass ratio). The efficient use of resources for above-ground space sequestration proved to be a parameter that quantitatively reflects competitiveness.

Summary • In the sensitive species Mimosa pudica electric signals arise when the leaves are stimulated by touching or wounding. Experiments reported here provide information about a photosynthetic response that results from heat-induced electrical signalling in leaves. • Electric potential measurements, combined with chlorophyll fluorescence, as well as gas exchange measurements showed that wounding evokes an electrical signal that travels rapidly into the neighbouring leaf pinna to eliminate the net-CO 2 uptake rate. At the same time the PSII quantum yield of electron transport is reduced from c . 0.6 to 0.2. Two-dimensional imaging analysis of the chlorophyll fluorescence signal revealed that the yield reduction spreads acropetally through the pinna and via the veins through the leaflets. •T o determine the speed of a chemical signal, a part of a pinna was exposed to 14 CO 2 . The remaining parts of the leaf were provided with label only when the translocation was extended to 12 h, indicating that a chemical signal is much too slow to account for the photosynthetic response after heat stimulation. • The results provide evidence for a role of the electrical signal in the regulation of photosynthesis because the high speed of the signal transduction rules out the involvement of a chemical signal, and the photosynthetic response occurs after the arrival of the electrical signal in the leaf pinna.

The occurrence of oxidative stress and oxidative damage in the halophyte Mesembryanthemum crystallinum during C 3 /crassulacean acid metabolism (CAM) transition evoked by salinity and high irradiance was investigated. The hypothesis tested was that CAM induction will effectively protect M. crystallinum against oxidative damage even under additionally applied oxidative stress via ozone (150 p.p.b.) exposure. Salinity (0.4 M NaCl), high irradiance and ozone induced the cytosolic CuZn-superoxide dismu- tase (SOD; EC 1.15.1.1) transcript abundance and activity indicating oxidative stress. However, ozone did not induce CAM. Contrary to the hypothesis, that ethylene may be involved in CAM induction, gassing with ethylene (200 p.p.b.) did not lead to CAM, but revealed that in M. crystallinum ethylene biosynthesis is induced by autostim- ulation. A putative fragment of the hitherto unknown nu- cleotide sequence of 1-aminocyclopropane-1-carboxylate oxidase (ACC; EC 4.4.1.14) was isolated and used as a specific probe against the ACC oxidase transcript to moni- tor oxidative damage. Ozone exposure increased the ACC oxidase expression only in plants with C 3 -photosynthesis which also showed severe necrosis and reduced photosyn- thetic activity, whereas plants in CAM mode (salt induced) remained unaffected. Natural stressors such as salinity and high irradiance induce CAM and up-regulate the ARS component CuZn-SOD very effectively to protect M. crys- tallinum from oxidative damage even when additional oxi- dative stress such as ozone is present.

A concept is presented for analysing and quantifying the competitiveness of woody plants. Efficiency ratios are defined that relate resource investments and gains to above- and belowground space sequestration. Such efficiencies are exemplified in comparisons between young and mature trees of European beech (Fagus sylvatica) and Norway spruce (Picea abies), resulting in remarkable consistencies across plant age and species. The demonstration extends to assessments of the inter- and intra-specific competition between young beech and spruce plants in a phytotron study, using CO2/O3 regimes as disturbants for examining the sensitivity of the efficiency ratios. In mixed plantations, the growth of spruce was enhanced and that of beech reduced as compared with the corresponding monocultures. This effect related to a high efficiency of spruce in aboveground space sequestration per resource investment, whereas the efficiency in carbon gain (return per unit of crown volume) was of minor importance. In contrast, beech displayed advantages in belowground space sequestration. However, competitiveness of young trees was hardly affected by the applied CO2/O3 treatments. The concept proves to be adequate for quantitatively comparing competitiveness across species, age and growth conditions. One further efficiency ratio, i.e., volume-related respiratory and transpiratory costs for sustaining functionality in resource sequestration, deserves attention in achieving a mechanistic understanding of competitive behaviour, in particular, in woody plants.

This study quantified the effect of soil warming on sap flow density ( Qs) of Pinus cembra L. at the treeline in the Central Tyrolean Alps. To enhance soil temperature we installed a transparent roof construction above the forest floor around six trees. Six other trees served as controls in the absence of any manipulation. Roofing enhanced growing season mean soil temperature by 1.6, 1.3 and 1.0 °C at 5, 10 and 20 cm soil depth, respectively, while soil water availability was not affected. Sap flow density (using Granier-type thermal dissipation probes) and environmental parameters were monitored throughout three growing seasons. During the first year of treatment, no warming effect was detected on Qs. However, soil warming caused Qs to increase signifi cantly by 11 and 19% above levels in control trees during the second and third year, respectively. This effect appeared to result from warming-induced root production, a reduction in viscosity and perhaps an increase also in root hydraulic conductivity. Hardly affected were leaf-level net CO2 uptake rate and conductance for water vapour, so that water-use efficiency stayed unchanged as confirmed by needle δ 13 C analysis. We conclude that tree water loss will increase with soil warming, which may alter the water balance within the treeline ecotone of the Central Austrian Alps in a future warming environment.

summary Young trees of European beech (Fagus sylvatica) acclimated for one growing season to ambient (c. 367 l ll ’" )o r elevated CO # levels (c. 660 l ll ’") were exposed during the subsequent year to combinations of the same CO # regimes and ambient or twice-ambient ozone (O $ ) levels (generated from the database of a rural site). By the end of June, before the development of macroscopic leaf injury, the raised O $ levels had not aected the light and dark reactions of photosynthesis. However, acclimation to elevated CO # had resulted in lowered chlorophyll and nitrogen concentrations, whereas photosynthetic performance, examined over a wide range of parameters from light and dark reactions, remained unchanged or showed only slight reductions (e.g. apparent electron transport rate, ETR; apparent quantum yield of CO # gas exchange, U CO# ; apparent carboxylation eciency, CE; and photosynthetic capacity at light and CO # saturation, PC). In August, after the appearance of leaf necroses, plants grown under ambient CO # and twice-ambient O $ conditions declined in both the photosynthetic light reactions (optimum electron quantum yield, F v }F m , non-photochemical energy quenching, NPQ, reduction state of Q A , apparent electron quantum yield, U PSI I, maximum electron transport rates) and the dark reactions as reflected by CE, U CO# , as well as the maximum CO # uptake rate (i.e. PC). CE, U CO# and PC were reduced by c. 75, 40 and 75%, respectively, relative to plants exposed to ambient CO # and O $ levels. By contrast, plants exposed to twice-ambient O $ and elevated CO # levels maintained a photosynthetic performance similar to individuals grown either under ambient CO # and ambient O $ , or elevated CO # and ambient O $ conditions. The long-term exposure to elevated CO # therefore tended to counteract adverse chronic eects of enhanced O $ levels on photosynthesis. Possible reasons for this compensatory eect in F. sylvatica are discussed.

The large majority of the Ca. 150 species of the neotropical shrub and tree genus Clusia have the potential to perform Crassulacean acid metabolism (CAM). They are either obligate CAM plants or C3/CAM intermediate plants. Only a very small percentage of the plants studied so far are apparently obligate C3 species. Among these was C. criuva, until recent laboratory studies showed that it may also have a certain CAM capacity under artificial stress conditions. Measurements of stomatal conductance (porometry) and chlorophyll fluorescence variables of C. criuva occurring along a transect from deep shade inside a gallery forest across the semi-shaded ecotone towards a cerrado and into the exposed cerrado itself in central Brazil now show that it can adapt its photosynthetic apparatus to effective performance of C3 photosynthesis under highly different photosynthetic photon flux densities. In addition, however, it does have a certain potential for CAM and cannot be considered as a strictly obligate C3 plant. Should a basic capacity for performing CAM be a general property of the genus, the quest for CAM traits in other remaining putatively obligate C3 species of the genus ought to be pursued.

Drought reduces the carbon (C) assimilation of trees and decouples aboveground from belowground carbon fluxes, but little is known about the response of drought-stressed trees to rewetting. This study aims to assess dynamics and patterns of C allocation in beech saplings under dry and rewetted soil conditions. In October 2010, 5-year-old beech saplings from a forest site were transplanted into 20 l pots. In 2011, the saplings were subjected to different levels of soil drought ranging from non-limiting water supply (control) to severe water limitation with soil water potentials of less than -1.5 MPa. As a physiologically relevant measure of drought, the cumulated soil water potential (i.e., drought stress dose (DSD)) was calculated for the growing season. In late August, the saplings were transferred into a climate chamber and pulse-labeled with (13)C-depleted CO2 (δ(13)C of -47‰). Isotopic signatures in leaf and soil respiration were repeatedly measured. Five days after soil rewetting, a second label was applied using 99 atom% (13)CO2. After another 12 days, the fate of assimilated C in each sapling was assessed by calculating the (13)C mass balance. Photosynthesis decreased by 60% in saplings under severe drought. The mean residence time (MRT) of recent assimilates in leaf respiration was more than three times longer than under non-limited conditions and was positively correlated to DSD. Also, the appearance of the label in soil respiration was delayed. Within 5 days after rewetting, photosynthesis, MRT of recent assimilates in leaf respiration and appearance of the label in soil respiration recovered fully. Despite the fast recovery, less label was recovered in the biomass of the previously drought-stressed plants, which also allocated less C to the root compartment (45 vs 64% in the control). We conclude that beech saplings quickly recover from extreme soil drought, although transitional after-effects prevail in C allocation, possibly due to repair-driven respiratory processes.

Diurnal measurements of chlorophyll fluorescence parameters and stomatal conductance to water vapour were carried out in five woody species and a bromeliad from two coastal vegetation types in Brazil, the rock outcrop of Pao de Acucar and the sand dunes of Macae. The environmental conditions of both study sites are characterized by high temperatures and light levels. The studied species comprised C3 plants with different degrees of stomatal closure during the day, overall daily stomatal conductance and a plant with a typical CAM pattern. Plants on Pao de Acucar exhibited only a small decline in potential quantum yield throughout the day. The non-photochemical quenching and the approximate photosynthetic electron transport rates were maximal during the peak of irradiance. In Macae, light response curves of fluorescence parameters in the CAM-tree Clusia hilariana showed a clear differentiation between phases III and IV of CAM. In phase III, decarboxylation of organic acids probably maintained high internal CO2 levels and there was only a small decrease in photochemical quenching with saturating light levels. In phase IV, the depletion of the organic acid pool and low stomatal conductances resulted in much lower levels of effective quantum yield and a substantial increase in both 1-qP and non-photochemical quenching with increasing light levels. This behaviour during phase IV in the CAM-tree C. hilariana was comparable to the performance of the C3-ClusiaC. aff parviflora at Pao de Acucar. It is seen that both the C3 species and the CAM tree occurring in the two coastal communities effectively adjust their photochemical efficiency to environmental conditions and especially to diurnal variations of stress.

Under continuous light the endogenous Crassulacean acid metabolism (CAM) rhythm of Kalanchoe daigremontiana Hamet et Perrier de la Bathie disappears at high (>29.0[deg]C) or low (

Changes in chlorophyll a fluorescence during the day and diurnal-changes of net CO2-exchange and organic acid contents were determined in two species of the genus Clusia during the dry season in Venezuela. The investigations included plants of the C3/CAM intermediate species Clusia minor and the C3 species C. multiflora growing at exposed and shaded sites. Both species showed a C3 pattern of net CO2-exchange at the exposed site. In the shade under extreme drought stress C. minor showed a weak expression of CAM without CO2-uptake during the afternoon (phase IV of CAM). C. multiflora growing in the shade exhibited a C3-pattern of net CO2-exchange and a small but significant nocturnal accumulation of citrate. Shaded plants of C. minor were able to double their light utilisation for electron transport and to reduce non-photochemical quenching during phase III compared to phase II of CAM. Furthermore, increase of electron transport rate through photosystem II in phase III of CAM is correlated to decarboxylation of malate. At the exposed site C. multiflora was less negatively affected by high PPFD than C. minor. This was shown by a lower reduction of potential electron quantum yield (Fv/Fm) and higher light utilisation of electron transport of C. multiflora compared to C. minor. At the exposed site C. minor did not make use of the CAM option to increase light utilisation of electron transport and to reduce non-photochemical quenching as did the plants growing in the shade.

Four different Clusia species showing C 3 photosynthesis (C. multiflora) or crassulacean acid metabolism (CAM) (C. rosea, an unidentified Clusia sp. and C. alata) co-occur in the same habitat in the northwest of Venezuela. The aim of this field study was to correlate the adaptive changes in chlorophyll a fluorescence with gas exchange patterns and deacidification of nocturnally accumulated organic acids. The results indicate that in C. rosea and Clusia sp. a lowering of the potential quantum yield, measured as F v /F m after 10 min of dark adaptation, is restricted to the early morning hours. With the onset of deacidification there was no further decline in F v /F m , although the irradiance upon the leaves remained high. For the C 3 -plant, C. multiflora, changes in F v /F m were correlated with changes in irradiance incident on the leaves before predarkening for 10 min for F v /F m measurements. The fast relaxation of F v /F m under low light indicates that C. multiflora was not suffering photoinhibitory damage. Effective quantum yield as determined by ΔF/F' m , was low under high irradiances for C. rosea and Clusia sp. and did not change significantly for C. rosea during phase III of CAM. During the time of low incident irradiance C. alata and C. multiflora reached an effective quantum yield that was close to the optimal quantum yield of electron transport. It is noteworthy that apparent electron transport rates through photosystem II (PS II) ((ΔF/F' m ) x PPFD) were highest for C. rosea, Clusia sp. and C. alata during phase III of CAM. Electron transport rates in C. multiflora were lower under high irradiance and did not differ from those found in phases II and IV of CAM. The possible role of CO 2 evolution from acid decarboxylation with respect to photosynthetic efficiency is discussed.

Samples of the Clusiaceae generaClusia, Oedematopus andDystovomita were collected at various sites and different altitudes in northern and south-western Venezuela. Analyses of stable isotopes of carbon and hydrogen and of leaf-nitrogen levels were performed on the dried samples. Correlations among these variables, i.e. carbon isotope discrimination (Δ), hydrogen isotope ratio (δD) and N-levels, and with altitude were assessed. In the samples, where values of Δ above 15‰ indicate predominant performance of C3 photosynthesis, there were slight tendencies of increasing Δ, δD and N-levels with increasing altitude and of increasing Δ with increasing N. Although these correlations taken separately were not statistically significant, they support each other and indicate increasing transpiration and increased leaf-nutrient supply at increasing altitude. Performance of crassulacean acid metabolism (CAM) in species ofClusia appears to be restricted to altitudes below 1500 m a.s.l. There was a significant negative correlation of Δ with altitude in the samples, where values of Δ below 10‰ indicated predominant performance of CAM. This suggests that phases II and IV of CAM are progressively suppressed towards the upper altitudinal limit of CAM inClusia in northern Venezuela. It is concluded that among the large number of environmental factors and combinations thereof, which determine the expression of CAM inClusia and trigger C3-CAM transitions in C3/CAM intermediate species, low availability of water is the most important.

Sandy plains are characteristic of the coastal region of Brazil. We investigated the diel patterns of changes in organic acid levels, leaf conductance and chlorophylla fluorescence for sun-exposed and shaded plants ofClusia hilariana, one of the dominant woody species in the sandy coastal plains of northern Rio de Janeiro state. Both exposed and shaded plants showed a typical CAM pattern with considerable diel oscillations in organic acid levels. The degradation of both malic and citric acids during the midday stomatal closure period could lead to potential CO2 fixation rates of 28 μmol m-2 s-1 in exposed leaves. Moreover, exposed leaves exhibited large increases in total non-photochemical quenching (qN) accompanied by a substantial decrease in effective quantum yield during the course of the day. However, these potential high rates of CO2 fixation and the increases inqn of exposed plants were not enough to maintain the primary electron acceptor of photosystem II (qA) in a low reduction state, similar to that of shaded plants. As a result, there was a moderate increase in the reduction state of qA throughout the day. Most of the decline in photochemical efficiency of exposed leaves ofC. hilariana was reversible, as evidenced by the high levels of pre-dawn potential quantum yields (Fv/Fm) and their rapid recovery after sunset. However, the depletion of the organic acid pool in the afternoon resulted in an accentuated subsequent drop in Fv/Fm, suggesting that prolonged periods of water stress accompanied by high irradiance levels may expose plants ofC. hilariana in unprotected habitats to the danger of photoinhibition.

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.

Combined delta(13)C and delta(18)O analyses of leaf material were used to infer changes in photosynthetic capacity (A(max)) and stomatal conductance (g(l)) in Fagus sylvatica and Picea abies trees growing under natural and controlled conditions. Correlation between g(l) and delta(18)O in leaf cellulose (delta(18)O(cel)) allowed us to apply a semi-quantitative model to infer g(l) from delta(18)O(cel) and also interpret variation in delta(13)C as reflecting variation in A(max). Extraction of leaf cellulose was necessary, because delta(18)O from leaf organic matter (delta(18)O(LOM)) and delta(18)O(cel) was not reliably correlated. In juvenile trees, the model predicted elevated carbon dioxide (CO(2)) to reduce A(max) in both species, whereas ozone (O(3)) only affected beech by reducing CO(2) uptake via lowered g(l). In adult trees, A(max) declined with decreasing light level as g(l) was unchanged. O(3) did not significantly affect isotopic signatures in leaves of adult trees, reflecting the higher O(3) susceptibility of juvenile trees under controlled conditions. The isotopic analysis compared favourably to the performance of leaf gas exchange, underlining that the semi-quantitative model approach provides a robust way to gather time-integrated information on photosynthetic performance of trees under multi-faced ecological scenarios, in particular when information needed for quantitative modelling is only scarcely available.

Root respiration has been shown to increase with temperature, but less is known about how this relationship is affected by the fungal partner in mycorrhizal root systems. In order to test respiratory temperature dependence, in particular Q (10) of mycorrhizal and non-mycorrhizal root systems, seedlings of PICEA ABIES (L.) Karst. (Norway spruce) were inoculated with the ectomycorrhizal fungus PILODERMA CROCEUM (Eriksson and Hjortstam, SR430; synonym: PILODERMA FALLAX: [Libert] Stalpers) and planted in soil respiration cuvettes (mycocosms). Temperature dependence of hyphal respiration in sterile cultures was determined and compared with respiration of mycorrhizal roots. Respiration rates of mycorrhizal and non-mycorrhizal root systems as well as sterile cultures were sensitive to temperature. Q (10) of mycorrhizal root systems of 3.0 +/- 0.1 was significantly higher than that of non-mycorrhizal systems (2.5 +/- 0.2). Q (10) of P. CROCEUM in sterile cultures (older than 2 months) was similar to that of mycorrhizal root systems, suggesting that mycorrhizae may have a large influence on the temperature sensitivity of roots in spite of their small biomass. Our results stress the importance of considering mycorrhization when modeling the temperature sensitivity of spruce roots.

The hypothesis that electric and hydraulic long-distance signals modify photosynthesis and stomatal aperture upon re-irrigation in intact drought-stressed plants was examined. Maize plants (Zea mays L.) were exposed to drought conditions by decreasing the soil water content to 40–50% of field capacity. The decrease in water content resulted in a decline in stomatal conductance to 50–60% of the level in well-watered plants. Re-irrigation of the plants initiated both hydraulic and electric signals, followed by a two-phase response of the net CO2 uptake rate and stomatal conductance of leaves. The transitional first phase (phase 1) is characterized by a rapid decrease in both levels. In the second phase (phase 2), both parameters gradually increase to levels above those of drought-stressed plants. Elimination of either the hydraulic signal by compensatory pressure application to the root system, or of the electric signal by cooling of the leaf blade gave evidence that the two signals (1) propagated independently from each other and (2) triggered the two-phase response in leaf gas exchange. The results provided evidence that the hydraulic signal initiated a hydropassive decrease in stomatal aperture and for the involvement of electric signals in the regulation of photosynthesis of drought-stressed plants.