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The widespread ascomycetous fungus Diplodia pinea is a latent, necrotrophic pathogen in Pinus species causing severe damages and world-wide economic losses. However, the interactions between pine hosts and virulent D. pinea are largely not understood. In the present study, systemic defence responses were investigated in non-inoculated, asymptomatic needles and roots of D. pinea infected saplings of two P. sylvestris provenances under controlled greenhouse conditions. Here, we show that D. pinea infection induced a multitude of systemic responses of the phytohormone profiles and metabolic traits. Shared systemic responses of both pine provenances in needles and roots included increased abscisic acid and jasmonic acid levels. Exclusively in the roots of both provenances, enhanced salicylic acid and reduced indole-3-acetic acid levels, structural biomass, and elevated activities of anti-oxidative enzymes were observed. Despite these similarities, the two pine provenances investigated different significantly in the systemic responses of both, phytohormone profiles and metabolic traits in needles and roots. However, the different systemic responses did not prevent subsequent destruction of non-inoculated needles, but rather prevented damage to the roots. Our results provide a detailed view on systemic defence mechanisms of pine hosts that are of particular significance for the selection of provenances with improved defence capacity., (© 2021 John Wiley & Sons Ltd.)

Nitric oxide (NO) is essential for plant growth and development, as well as interactions with abiotic and biotic environments. Its importance for multiple functions in plants means that tight regulation of NO concentrations is required. This is of particular significance in roots, where NO signalling is involved in processes, such as root growth, lateral root formation, nutrient acquisition, heavy metal homeostasis, symbiotic nitrogen fixation and root-mycorrhizal fungi interactions. The NO signal can also be produced in high levels by microbial processes in the rhizosphere, further impacting root processes. To explore these interesting interactions, in the present review, we firstly summarize current knowledge of physiological processes of NO production and consumption in roots and, thereafter, of processes involved in NO homeostasis in root cells with particular emphasis on root growth, development, nutrient acquisition, environmental stresses and organismic interactions., (© 2020 John Wiley & Sons Ltd.)

Uptake, translocation, detoxification, and sequestration of heavy metals (HMs) are key processes in plants to deal with excess amounts of HM. Under natural conditions, plant roots often establish ecto- and/or arbuscular-mycorrhizae with their fungal partners, thereby altering HM accumulation in host plants. This review considers the progress in understanding the physiological and molecular mechanisms involved in HM accumulation in nonmycorrhizal versus mycorrhizal plants. In nonmycorrhizal plants, HM ions in the cells can be detoxified with the aid of several chelators. Furthermore, HMs can be sequestered in cell walls, vacuoles, and the Golgi apparatus of plants. The uptake and translocation of HMs are mediated by members of ZIPs, NRAMPs, and HMAs, and HM detoxification and sequestration are mainly modulated by members of ABCs and MTPs in nonmycorrhizal plants. Mycorrhizal-induced changes in HM accumulation in plants are mainly due to HM sequestration by fungal partners and improvements in the nutritional and antioxidative status of host plants. Furthermore, mycorrhizal fungi can trigger the differential expression of genes involved in HM accumulation in both partners. Understanding the molecular mechanisms that underlie HM accumulation in mycorrhizal plants is crucial for the utilization of fungi and their host plants to remediate HM-contaminated soils., (© 2018 John Wiley & Sons Ltd.)

We used instantaneous temperature responses of CO 2 -respiration to explore temperature acclimation dynamics for Eucalyptus grandis grown with differing nitrogen supply. A reduction in ambient temperature from 23 to 19 °C reduced light-saturated photosynthesis by 25% but increased respiratory capacity by 30%. Changes in respiratory capacity were not reversed after temperatures were subsequently increased to 27 °C. Temperature sensitivity of respiration measured at prevalent ambient temperature varied little between temperature treatments but was significantly reduced from ~105 kJ mol -1 when supply of N was weak, to ~70 kJ mol -1 when it was strong. Temperature sensitivity of respiration measured across a broader temperature range (20-40 °C) could be fully described by 2 exponent parameters of an Arrhenius-type model (i.e., activation energy of respiration at low reference temperature and a parameter describing the temperature dependence of activation energy). These 2 parameters were strongly correlated, statistically explaining 74% of observed variation. Residual variation was linked to treatment-induced changes in respiration at low reference temperature or respiratory capacity. Leaf contents of starch and soluble sugars suggest that respiratory capacity varies with source-sink imbalances in carbohydrate utilization, which in combination with shifts in carbon-flux mode, serve to maintain homeostasis of respiratory temperature sensitivity at prevalent growth temperature., (© 2018 John Wiley & Sons Ltd.)

Phytopathogenic fungi infections induce plant defence responses that mediate changes in metabolic and signalling processes with severe consequences for plant growth and development. Sphaeropsis tip blight, induced by the endophytic fungus Sphaeropsis sapinea that spreads from stem tissues to the needles, is the most widespread disease of conifer forests causing dramatic economic losses. However, metabolic consequences of this disease on bark and wood tissues of its host are largely unexplored. Here, we show that diseased host pines experience tissue dehydration in both bark and wood. Increased cytokinin and declined indole-3-acetic acid levels were observed in both tissues and increased jasmonic acid and abscisic acid levels exclusively in the wood. Increased lignin contents at the expense of holo-cellulose with declined structural biomass of the wood reflect cell wall fortification by S. sapinea infection. These changes are consistent with H 2 O 2 accumulation in the wood, required for lignin polymerization. Accumulation of H 2 O 2 was associated with more oxidized redox states of glutathione and ascorbate pools. These findings indicate that S. sapinea affects both phytohormone signalling and the antioxidative defence system in stem tissues of its pine host during the infection process., (© 2017 John Wiley & Sons Ltd.)

Here, we characterized nitrogen (N) uptake of beech (Fagus sylvatica) and their associated ectomycorrhizal (EM) communities from NH 4 + and NO 3 - . We hypothesized that a proportional fraction of ectomycorrhizal N uptake is transferred to the host, thereby resulting in the same uptake patterns of plants and their associated mycorrhizal communities. 15 N uptake was studied under various field conditions after short-term and long-term exposure to a pulse of equimolar NH 4 + and NO 3 - concentrations, where one compound was replaced by 15 N. In native EM assemblages, long-term and short-term 15 N uptake from NH 4 + was higher than that from NO 3 - , regardless of season, water availability and site exposure, whereas in beech long-term 15 N uptake from NO 3 - was higher than that from NH 4 + . The transfer rates from the EM to beech were lower for 15 N from NH 4 + than from NO 3 - . 15 N content in EM was correlated with 15 N uptake of the host for 15 NH 4 + , but not for 15 NO 3 - -derived N. These findings suggest stronger control of the EM assemblage on N provision to the host from NH 4 + than from NO 3 - . Different host and EM accumulation patterns for inorganic N will result in complementary resource use, which might be advantageous in forest ecosystems with limited N availability., (© 2016 John Wiley & Sons Ltd.)

One major effect of global climate change will be altered precipitation patterns in many regions of the world. This will cause a higher probability of long-term waterlogging in winter/spring and flash floods in summer because of extreme rainfall events. Particularly, trees not adapted at their natural site to such waterlogging stress can be impaired. Despite the enormous economic, ecological and social importance of forest ecosystems, the effect of waterlogging on trees is far less understood than the effect on many crops or the model plant Arabidopsis. There is only a handful of studies available investigating the transcriptome and metabolome of waterlogged trees. Main physiological responses of trees to waterlogging include the stimulation of fermentative pathways and an accelerated glycolytic flux. Many energy-consuming, anabolic processes are slowed down to overcome the energy crisis mediated by waterlogging. A crucial feature of waterlogging tolerance is the steady supply of glycolysis with carbohydrates, particularly in the roots; stress-sensitive trees fail to maintain sufficient carbohydrate availability resulting in the dieback of the stressed tissues. The present review summarizes physiological and molecular features of waterlogging tolerance of trees; the focus is on carbon metabolism in both, leaves and roots of trees., (© 2014 John Wiley & Sons Ltd.)

Sulphate uptake and its distribution within plants depend on the activity of different sulphate transporters (SULTR). In long-living deciduous plants such as trees, seasonal changes of spatial patterns add another layer of complexity to the question of how the interplay of different transporters adjusts S distribution within the plant to environmental changes. Poplar is an excellent model to address this question because its S metabolism is already well characterized. In the present study, the importance of SULTRs for seasonal sulphate storage and mobilization was examined in the wood of poplar (Populus tremula × P. alba) by analysing their gene expression in relation to sulphate contents in wood and xylem sap. According to these results, possible functions of the respective SULTRs for seasonal sulphate storage and mobilization in the wood are suggested. Together, the present results complement the previously published model for seasonal sulphate circulation between leaves and bark and provide information for future mechanistic modelling of whole tree sulphate fluxes., (© 2012 John Wiley & Sons Ltd.)

We investigated the interaction of rhizospheric nitric oxide (NO) concentration (i.e. low, ambient or high) and soil nitrogen (N) availability (i.e. low or high) with organic and inorganic N uptake by fine roots of Pinus sylvestris L. seedlings by (15) N feeding experiments under controlled conditions. N metabolites in fine roots were analysed to link N uptake to N nutrition. NO affected N uptake depending on N source and soil N availability. The suppression of nitrate uptake in the presence of ammonium and glutamine was overruled by high NO. The effects of NO on N uptake with increasing N availability showed different patterns: (1) increasing N uptake regardless of NO concentration (i.e. ammonium); (2) increasing N uptake only with high NO concentration (i.e. nitrate and arginine); and (3) decreasing N uptake (i.e. glutamine). At low N availability and high NO nitrate accumulated in the roots indicating insufficient substrates for nitrate reduction or its storage in root vacuoles. Individual amino acid concentrations were negatively affected with increasing NO (i.e. asparagine and glutamine with low N availability, serine and proline with high N availability). In conclusion, this study provides first evidence that NO affects N uptake and metabolism in a conifer., (© 2012 Blackwell Publishing Ltd.)

In the present study, the significance of sulfite oxidase (SO) for sulfite detoxification and sulfur assimilation was investigated. In response to sulfur dioxide (SO(2)) exposure, a remarkable expansion of sulfate and a significant increase of GSH pool were observed in wild-type and SO-overexpressing Arabidopsis. These metabolic changes were connected with a negative feedback inhibition of adenosine 5'-phosphosulfate reductase (APR), but no alterations in gas exchange parameters or visible symptoms of injury. However, Arabidopsis SO-KO mutants were consistently negatively affected upon 600 nL L(-1) SO(2) exposure for 60 h and showed phenotypical symptoms of injury with small necrotic spots on the leaves. The mean g(H2O) was reduced by about 60% over the fumigation period, accompanied by a reduction of net CO(2) assimilation and SO(2) uptake of about 50 and 35%. Moreover, sulfur metabolism was completely distorted. Whereas sulfate pool was kept constant, thiol-levels strongly increased. This demonstrates that SO should be the only protagonist for back-oxidizing and detoxification of sulfite. Based on these results, it is suggested that co-regulation of SO and APR controls sulfate assimilation pathway and stabilizes sulfite distribution into organic sulfur compounds. In conclusion, a sulfate-sulfite cycle driven by APR and SO can be postulated for fine-tuning of sulfur distribution that is additionally used for sulfite detoxification, when plants are exposed to atmospheric SO(2)., (© 2011 Blackwell Publishing Ltd.)

The analysis of delta(13)C and delta(18)O in tree-ring archives offers retrospective insights into environmental conditions and ecophysiological processes. While photosynthetic carbon isotope discrimination and evaporative oxygen isotope enrichment are well understood, we lack information on how the isotope signal is altered by downstream metabolic processes. In Pinus sylvestris, we traced the isotopic signals from their origin in the leaf water (delta(18)O) or the newly assimilated carbon (delta(13)C), via phloem sugars to the tree-ring, over a time-scale that ranges from hours to a growing season. Seasonally, variable (13)C enrichment of sugars related to phloem loading and transport did lead to uncoupling between delta(13)C in the tree-ring, and the c(i)/c(a) ratio at the leaf level. In contrast, the oxygen isotope signal was transferred from the leaf water to the tree-ring with an expected enrichment of 27 per thousand, with time-lags of approximately 2 weeks and with a 40% exchange between organic oxygen and xylem water oxygen during cellulose synthesis. This integrated overview of the fate of carbon and oxygen isotope signals within the model tree species P. sylvestris provides a novel physiological basis for the interpretation of delta(13)C and delta(18)O in tree-ring ecology.

Salinity represents an increasing environmental problem in managed ecosystems. Populus spp. is widely used for wood production by short-rotation forestry in fertilized plantations and can be grown on saline soil. Because N fertilization plays an important role in salt tolerance, we analysed Grey poplar (Populus tremula x alba, syn. Populus canescens) grown with either 1 mM nitrate or ammonium subjected to moderate 75 mM NaCl. The impact of N nutrition on amelioration of salt tolerance was analysed on different levels of N metabolism such as N uptake, assimilation and N (total N, proteins and amino compounds) accumulation. Na concentration increased in all tissues over time of salt exposure. The N nutrition-dependent effects of salt exposure were more intensive in roots than in leaves. Application of salt reduced root increment as well as stem height increase and, at the same time, increased the concentration of total amino compounds more intensively in roots of ammonium-fed plants. In leaves, salt treatment increased concentrations of total N more intensively in nitrate-fed plants and concentrations of amino compounds independently of N nutrition. The major changes in N metabolism of Grey poplar exposed to moderate salt concentrations were detected in the significant increase of amino acid concentrations. The present results indicate that N metabolism of Grey poplar exposed to salt performed better when the plants were fed with nitrate instead of ammonium as sole N source. Therefore, nitrate fertilization of poplar plantations grown on saline soil should be preferred.

Understanding ecosystem water fluxes has gained increasing attention, as climate scenarios predict a drier environment for many parts of the world. Evaporative enrichment of (18)O (Delta(18)O) of leaf water and subsequent enrichment of plant organic matter can be used to characterize environmental and physiological factors that control evaporation, based on a recently established mechanistic model. In a Pinus sylvestris forest, we measured the dynamics of oxygen isotopic composition (delta(18)O) every 6 h for 4 d in atmospheric water vapour, xylem sap, leaf water and water-soluble organic matter in current (N) and previous year (N-1) needles, phloem sap, together with leaf gas exchange for pooled N and N-1 needles, and relevant micrometeorological variables. Leaf water delta(18)O showed strong diel periodicity, while delta(18)O in atmospheric water vapour and in xylem sap showed little variation. The Delta(18)O was consistently lower for N than for N-1 needles, possibly related to phenological stage. Modelled leaf water Delta(18)O showed good agreement with measured values when applying a non-steady state evaporative enrichment model including a Péclet effect. We determined the time lags between delta(18)O signals from leaf water to water-soluble foliar organic matter and to phloem sap at different locations down the trunk, which clearly demonstrated the relevance of considering these time-lag effects for carbon transport, source-sink and carbon flux partitioning studies.

Sulphur dioxide (SO(2)) is known as a strongly damaging air pollutant. After conversion to sulphite in aqueous solution, it becomes a strong nucleophilic agent that attacks numerous compounds in the cell. Therefore, plants have developed a mechanism to control sulphite levels. Recently, we have cloned and characterized the enzyme sulphite oxidase (SO) from Arabidopsis thaliana. Yet, its physiological role remained unclear. Here, we describe results demonstrating that SO is essential for detoxifying excessive amounts of sulphite in the cell which is important for the survival of the plant. T-DNA-tagged A. thaliana plants lacking the enzyme showed a decrease in vitality during SO(2) fumigation and a change in their S-metabolites. The same was found with RNA-interference (RNAi) plants that were generated for tobacco. On the contrary, over-expression of SO helped the plant to survive SO(2) concentrations that are detrimental for non-transformed wild-type (WT) plants, as was shown with poplar plants which are known to be particularly sensitive to SO(2). Fumigation induced the expression of the enzyme as demonstrated by promoter-reporter gene fusion, by immunoblot analysis of SO-protein and by induction of enzyme activity. This implies that SO, as an otherwise constitutively expressed protein, is under additional control by SO(2) in the environment.

This study investigated the influence of meteorological, pedospheric and physiological factors on the water relations of Scots pine, as characterized by the origin of water taken up, by xylem transport as well as by carbon isotope discrimination (Delta13C) and oxygen isotope enrichment (Delta18O) of newly assimilated organic matter. For more than 1 year, we quantified delta2H and delta18O of potential water sources and xylem water as well as Delta13C and Delta18O in twig and trunk phloem organic matter biweekly, and related these values to continuously measured or modelled meteorological parameters, soil water content, stand transpiration (ST) and canopy stomatal conductance (G(s)). During the growing season, delta18O and delta2H of xylem water were generally in a range comparable to soil water from a depth of 2-20 cm. Long residence time of water in the tracheids uncoupled the isotopic signals of xylem and soil water in winter. Delta18O but not Delta13C in phloem organic matter was directly indicative of recent environmental conditions during the whole year. Delta18O could be described applying a model that included 18O fractionation associated with water exchange between leaf and atmosphere, and with the production of organic matter as well as the influence of transpiration. Phloem Delta13C was assumed to be concertedly influenced by G(s) and photosynthetically active radiation (PAR) (as a proxy for photosynthetic capacity). We conclude that isotope signatures can be used as effective tools (1) to characterize the seasonal dynamics in source and xylem water, and (2) to assess environmental effects on transpiration and G(s) of Scots pine, thus helping to understand and predict potential impacts of climate change on trees and forest ecosystems.

This study investigated the influence of climate on the carbon isotopic composition (sigma13C) and oxygen isotopic enrichment (delta18O) above the source water of different organic matter pools in European beech. In July and September 2002, sigma13C and delta18O were determined in phloem carbohydrates and in bulk foliage of adult beech trees along a transect from central Germany to southern France, where beech reaches its southernmost distributional limit. The data were related to meteorological and physiological parameters. The climate along the transect stretches from temperate [subcontinental (SC)] to submediterranean (SM). Both sigma13Cleaf and delta18Oleaf were representative of site-specific long-term environmental conditions. sigma13C of leaves collected in September was indicative of stomatal conductance, vapour pressure deficit (VPD) and radiation availability of the current growing season. delta18O was mainly correlated to mean growing season relative humidity (RH) and VPD. In contrast to the leaves, sigma13Cphloem varied considerably between July and September and was well correlated with canopy stomatal conductance (Gs) in a 2 d integral prior to phloem sampling. The relationship between sigma13C and delta18O in both leaves and phloem sap points, however, to a combined influence of stomatal conductance and photosynthetic capacity on the variation of sigma13C along the transect. delta18Ophloem could be described by applying a model that included 18O fractionation associated with water exchange between the leaf and the atmosphere and with the production of organic matter. Hence, isotope signatures can be used as effective tools to assess the water balance of beech, and thus, help predict the effects of climatic change on one of the ecologically and economically most important tree species in Central Europe.

The present study shows for the first time the influence of exogenously applied amino acids and cytokinin on the physiological and molecular aspects of N metabolism in poplar trees. In a short-term feeding experiment, glutamine or trans-zeatin riboside (tZR) was added directly to the nutrient solution. NO3- net uptake declined significantly in response to both treatments. Feeding with glutamine brought about an increase in concentrations of different amino compounds in the roots (glutamine, glutamate, alanine, gamma-amino butyric acid (GABA) and NH4+, which negatively correlated with the net NO3- uptake. The plants showed a reduction of cytosolic glutamine synthetase 1 (GS1) transcript level in the roots. In addition, glutamine feeding changed the root-to-shoot distribution on N assimilation in favour of the leaves and plant internal N cycling. tZR treatment resulted in expansion of zeatin-type (Z-type) cytokinins in the roots and increased nitrate reductase (NR)-mRNA level. The results indicate that both particular amino acids and active cytokinins are involved in the feedback regulation of N uptake and metabolism in poplar. We propose that inhibition of N uptake by cytokinins in poplar is more complex than that mediated by amino compounds, and other effectors are involved in this regulation.

Beech seedlings from 11 German climatic provenances were exposed to a realistically timed drought treatment in a greenhouse experiment. The stable isotope composition of carbon (C) and nitrogen (N) was analysed in pooled bulk material of roots, stems and leaves, as well as in the aqueous extracts and starch fractions. The delta 13C values increased in bulk samples (BS) of roots, stems and leaves by drought, although no leaf growth occurred during the experimental period. A clear drought effect on delta 13C in aqueous extracts was detected in leaves. In aqueous extracts of stems and roots as well as in starch fractions of all organs, abundance of delta 13C also tended to be increased by drought, but this effect was not statistically significant. For both delta 13C and delta 15N, enrichment was observed from the site of uptake/ source to the site of use/sink. A gradient for delta 13C in all fractions from leaves (-29.49, -28.89 and -27.85 per thousand) to stems (-28.81, -27.48 and -26.98 per thousand) and to roots (-27.60, -26.37 and -26.48 per thousand) was detected in BS, aqueous extracts and starch, respectively. An opposite gradient for delta 15N was found in BS: 1.59 per thousand, 1.84 per thousand and 3.05 per thousand in roots, stems and leaves, respectively. delta 15N was neither affected by drought in the BS nor in aqueous extracts, but an effect of provenance was observed. Particularly in roots and stems, drought-sensitive provenances showed the strongest shifts in delta 13C induced by drought and the lowest delta 15N values. In the present experiment, delta 13C values were more affected by the environmental factor drought, while delta 15N values were more affected by the genetic factor provenance.

The level of glutathione (GSH) in plants is important in defence reactions against biotic and abiotic stresses and can place considerable demand of the sulphur assimilation pathway. Enzymes involved in sulphur assimilation and GSH metabolism are not evenly distributed between different subcellular compartments or between different cell types in leaves or roots; however, there is little information on the effect that such asymmetries have on the actual GSH concentration in each compartment or cell type. In the present study in situ labelling with monochlorobimane (MCB) in combination with confocal laser scanning microscopy was used to quantify GSH in each of the main cell types of poplar leaves from fluorescence of the GSB conjugate formed. Comparison of results from the in situ approach with total GSH levels measured in vitro by high-performance liquid chromatography suggested that only the cytosolic GSH pool was labelled using this approach. This suggests that an appropriate GST was not present within the chloroplasts to catalyse the conjugation reaction and that chloroplastic GSH does not rapidly exchange with the cytoplasmic pool under the conditions of the assay. Cytosolic GSH levels were between 0.2 and 0.3 mm for both photosynthetic and non-photosynthetic (epidermal) cell types in wild-type poplar leaves. Cytosolic levels increased by around two-fold in transgenic poplars over-expressing bacterial gamma-glutamylsynthetase (gamma-ECS) in the cytosol of all cell types, but there was no concomitant increase in the chloroplastic GSH pool.