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IntroductionGroundcover management plays a crucial role in improving water retention and soil nutrition in orchard systems, thereby preventing environmental constrains by non-point source pollution. However, effectiveness of groundcover management in citrus orchards developed on sloping farmland with eroded purple soil has not been studied in detail. In particular, information on the soil nutrient losses, e.g., nitrogen (N) and phosphorus (P), through interflow and its effects on growth and nutrition of citrus plants has not been reported.MethodsThe present study evaluated the effects of different cover crops, i.e., Lolium perenne L. (Lolium), Vicia villosa Roth (Vicia) and Orychophragmus violaceus (Ory), on nutrition and growth of citrus trees as well as water, soil and nutrient retention in an orchard developed in sloping farmland during two consecutive years.Results and discussionThe results show that the groundcover species Lolium and Vicia mediated nursing effects on nutrition and growth of citrus trees. These nursing effects included enhanced foliar levels of carbon(C), N and P as well as increased tree height, stem diameter, and crown width. Groundcover management generally reduced the annual surface runoff, interflow, soil loss, total N loss and total P loss. Among the cover crop species studied, Lolium and Vicia were overall more efficient than Ory in this context. Lolium reduced the average annual total loss of N and P by 42.53% and 49.23%, respectively, compared with clean tillage. The estimated annual reduction potentials of soil, N and P losses in Southwestern China were 16.3, 3.4 and 8.5 million tons yr-1, respectively. Obviously, Lolium and Vicia provide highly beneficial ground coverage on sloping farmland and, thus, can be used for future sustainable development of citrus orchards.

Drought stress is a major environmental factor limiting citrus productivity. Still, differences in drought sensitivity between citrus hybrids of different maturation periods have so far not been reported. Here, we selected a medium-maturing (Fertile orange: FO (Citrus reticulata cv. Fertile orange) and a late-maturing citrus hybrid (Newhall Navel orange: NO (Citrus sinensis Osbeck cv. Newhall) and determined the physiological and biochemical traits of leaves, roots, wood and bark. Our results showed that drought significantly decreased net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (Tr) of citrus leaves. Oxidative stress upon drought was indicated by enhanced foliar malondialdehyde (MDA) and hydrogen peroxide contents, as well as a stimulation of the anti-oxidative system. This stimulation included the contents of dehydroascorbic acid (DHA), glutathione (GSH) and oxidized glutathione (GSSG) in leaves, roots, wood and bark, as well as activities of antioxidative enzymes of glutathione reductase (GR), dehydroascorbate reductase (DHAR), superoxide dismutase (SOD) and peroxidase (POD). The late maturing NO hybrid not only showed better general physiological performance as indicated by increased Pn in leaves, but also higher biochemical ROS scavenging and osmotic capacity as indicated by increased ascorbic acids (ASA), DHA, and proline contents, as well as activities of enzymes of SOD, POD, ASA/DHA and GSH/GSSG ratios in the investigated tissues compared to the FO hybrid under drought and control conditions. Analysis of molecular mechanisms of signaling, regulatory and functional genes expression are suggested for future studies to elucidate the complex interplay of molecular, biochemical and physiological responses of citrus hybrids to drought.

Top grafting is an efficient and practical technique for the renewal and rejuvenation of citrus trees in old orchards. However, root death after top grafting restricts plant growth and canopy reconstruction. Thus, applications of rooting promotion substances before citrus top grafting may increase the amount and activity of roots, thereby enhancing top-grafted plant performance. To test this assumption, four rooting promotion substances, i.e., rooting promotion powder, biochar, organic fertilizer, and potassium fulvic acid, were applied before top grafting, and the effects on biometric and physiological parameters were analyzed after top grafting. The results showed that the application of all rooting promotion substances before top grafting has a positive effect on growth and mineral nutrient acquisition, as well as on foliar C and N assimilates and the activity of anti-oxidative enzymes of top-grafted plants. Rooting promotion powder and biochar had the best effect on top-grafted tree performance in the short term. In conclusion, pre-grafting root promotion reduced root damage, enhanced nutrient acquisition, and improved the physiological performance of top-grafted plants. Therefore, this approach can play a crucial role in accelerating canopy reconstruction in old citrus orchards and in improving citrus plant development.

Abstract Date palm (Phoenix dactylifera L.) is able to grow and complete its life cycle while being rooted in highly saline soils. Which of the many well‐known salt‐tolerance strategies are combined to fine‐tune this remarkable resilience is unknown. The precise location, whether in the shoot or the root, where these strategies are employed remains uncertain, leaving us unaware of how the various known salt‐tolerance mechanisms are integrated to fine‐tune this remarkable resilience. To address this shortcoming, we exposed date palm to a salt stress dose equivalent to seawater for up to 4 weeks and applied integrative multi‐omics analyses followed by targeted metabolomics, hormone, and ion analyses. Integration of proteomic into transcriptomic data allowed a view beyond simple correlation, revealing a remarkably high degree of convergence between gene expression and protein abundance. This sheds a clear light on the acclimatization mechanisms employed, which depend on reprogramming of protein biosynthesis. For growth in highly saline habitats, date palm effectively combines various salt‐tolerance mechanisms found in both halophytes and glycophytes: “avoidance” by efficient sodium and chloride exclusion at the roots, and “acclimation” by osmotic adjustment, reactive oxygen species scavenging in leaves, and remodeling of the ribosome‐associated proteome in salt‐exposed root cells. Combined efficiently as in P. dactylifera L., these sets of mechanisms seem to explain the palm's excellent salt stress tolerance.

Abstract Background The cactus family (Cactaceae) has been reported to have evolved a minimal photosynthetic plastome size, with the loss of inverted-repeat (IR) regions and NDH gene suites. However, there are very limited genomic data on the family, especially Cereoideae, the largest subfamily of cacti. Results In the present study, we assembled and annotated 35 plastomes, 33 of which were representatives of Cereoideae, alongside 2 previously published plastomes. We analyzed the organelle genomes of 35 genera in the subfamily. These plastomes have variations rarely observed in those of other angiosperms, including size differences (with ~ 30 kb between the shortest and longest), dramatic dynamic changes in IR boundaries, frequent plastome inversions, and rearrangements. These results suggested that cacti have the most complex plastome evolution among angiosperms. Conclusion These results provide unique insight into the dynamic evolutionary history of Cereoideae plastomes and refine current knowledge of the relationships within the subfamily.

Deforestation is one of the most serious environmental problems facing humankind. It continues to escalate rapidly across many regions of the world, thereby deteriorating the forest soil quality. This has prompted a large number of field-based studies aimed at understanding the impacts of deforestation on soil properties. However, the lack of comprehensive meta-analyses that utilized these studies has limited our deeper understanding of how different soil properties, including the soil organic carbon (SOC) pool, respond to deforestation. To address this critical knowledge gap, we conducted a meta-analysis of 144 studies to explore the impacts of deforestation on soil chemical, physical, and biological properties, with special emphasis on the long-term changes in SOC, such as concentrations, stocks, and sequestration. The results revealed that deforestation significantly decreased soil organic matter, electrical conductivity, and base saturation by 52%, 50%, and 98%, respectively. While deforestation increased soil total nitrogen content and decreased available phosphorus content by 51% and 99%, respectively, it resulted in slight decreases in some chemical properties, including soil pH (1%) and base cations (1%–13%). Deforestation significantly increased bulk density by 27% and soil erosion by 47%, but significantly decreased soil aggregate stability by 39% and saturated hydraulic conductivity by 63%. Soil microbial biomass C and N concentrations and enzyme activities were significantly decreased as a consequence of deforestation. Soil biological properties were much more affected by deforestation than soil physical and chemical properties. Regarding the SOC, the land use conversion from forest to pasture significantly increased SOC concentrations, stocks, and sequestration rates (11%–13%), whereas the land use conversions from forest to both plantation and cropland significantly decreased SOC concentrations, stocks, and sequestration rates (10%–43%). This observed decline in SOC accumulations decreased with increasing years after deforestation. The SOC dynamics following deforestation were predominantly regulated by microbial biomass concentrations, dehydrogenase activity, soil erosion, saturated hydraulic conductivity, aggregate stability, as well as concentrations of total organic carbon, total nitrogen, total phosphorus and organic matter. The present meta-analytical study provides compelling evidence that deforestation can induce profound changes in soil characteristics, including soil C contents, and has significant implications for soil health sustainability and climate change mitigation.

The plant microbiota can affect plant health and fitness by promoting methylmercury (MeHg) production in paddy soil. Although most well-known mercury (Hg) methylators are observed in the soil, it remains unclear how rice rhizosphere assemblages alter MeHg production. Here, we used network analyses of microbial diversity to identify bulk soil (BS), rhizosphere (RS) and root bacterial networks during rice development at Hg gradients. Hg gradients greatly impacted the niche-sharing of taxa significantly relating to MeHg/THg, while plant development had little effect. In RS networks, Hg gradients increased the proportion of MeHg-related nodes in total nodes from 37.88% to 45.76%, but plant development enhanced from 48.59% to 50.41%. The module hub and connector in RS networks included taxa positively (Nitrososphaeracea, Vicinamibacteraceae and Oxalobacteraceae) and negatively (Gracilibacteraceae) correlating with MeHg/THg at the blooming stage. In BS networks, Deinococcaceae and Paludibacteraceae were positively related to MeHg/THg, and constituted the connector at the reviving stage and the module hub at the blooming stage. Soil with an Hg concentration of 30 mg kg−1 increased the complexity and connectivity of root microbial networks, although microbial community structure in roots was less affected by Hg gradients and plant development. As most frequent connector in root microbial networks, Desulfovibrionaceae did not significantly correlate with MeHg/THg, but was likely to play an important role in the response to Hg stress.

Drought-sensitive European beech forests are increasingly challenged by climate change. Admixing other, preferably more deep-rooting, tree species has been proposed to increase the resilience of beech forests to drought. This diversification of beech forests might also affect soil organic carbon (SOC) and total nitrogen (TN) stocks that are relevant for a wide range of soil functions and ecosystem services, such as water and nutrient retention, filter functions and erosion control. Since information of these effects is scattered, our aim was to synthesize results from studies that compared SOC/TN stocks of beech monocultures with those of beech stands mixed with other tree species as well as monocultures of other tree species. We conducted a meta-analysis including 38 studies with 203, 220, and 160 observations for forest floor (i.e., the organic surface layer), mineral soil (0.5 m depth) and the total soil profile, respectively. Monoculture conifer stands had higher SOC stocks compared to monoculture beech in general, especially in the forest floor (up to 200% in larch forests). In contrast, other broadleaved tree species (oak, ash, lime, maple, hornbeam) showed lower SOC stocks in the forest floor compared to beech, with little impact on total SOC stocks. Comparing mixed beech-conifer stands (average mixing ratio with regard to number of trees 50:50) with beech monocultures revealed significantly higher total SOC stocks of around 9% and a smaller increase in TN stocks of around 4%. This equaled a SOC accrual of 0.1 Mg ha−1 yr−1. In contrast, mixed beech-broadleaved stands did not show significant differences in total SOC stocks. Conifer admixture effects on beech forest SOC were of additive nature. Admixing other tree species to beech monoculture stands was most effective to increase SOC stocks on low carbon soils with a sandy texture and nitrogen limitation (i.e., a high C/N ratio and low nitrogen deposition). We conclude that, with targeted admixture measures of coniferous species, an increase in SOC stocks in beech forests can be achieved as part of the necessary adaptation of beech forests to climate change.

This is a novel study about responses of leaf photosynthetic traits and plant mercury (Hg) accumulation of rice grown in Hg polluted soils to elevated CO2 (ECO2). The aim of this study was to provide basic information on the acclimation capacity of photosynthesis and Hg accumulation in rice grown in Hg polluted soil under ECO2 at day, night, and full day. For this purpose, we analyzed leaf photosynthetic traits of rice at flowering and grain filling. In addition, chlorophyll content, soluble sugar and Malondialdehyde (MDA) of rice leaves were measured at flowering. Seed yield, ear number, grain number per ear, 1000-grain weight, total mercury (THg) and methylmercury (MeHg) contents were determined after harvest. Our results showed that Hg polluted soil and ECO2 had no significant effect on leaf chlorophyll content and leaf mass per area (LMA) in rice. The contents of soluble sugar and MDA in leaves increased significantly under ECO2. Mercury polluted soil treatment significantly reduced the light saturated CO2 assimilation rate (Asat) of rice leaves only at flowering, but not at grain filling. Night ECO2 greatly improved rice leaf water use efficiency (WUE). ECO2 greatly increased seed yield and ear number. In addition, ECO2 did not affect THg accumulation in rice organs, but ECO2 and Hg treatment had a significant interaction on MeHg in seeds, husks and roots.

Aleppo pine (Pinus halepensis) is widely distributed in the Mediterranean region and in other areas of the world, where it has been introduced due to its adaptive capacity to xerothermic conditions. The giant pine scale Marchalina hellenica often infests Aleppo pine, as well as other pines, in several southeastern European countries, causing pine declines. When combined with the expected intensified heat and drought events in eastern Mediterranean, the impact of this biotic parameter on the host pines may be exacerbated. The importance of understanding the defense mechanisms of Aleppo pine is emphasized by the recent invasion of the pine scale in new regions, like Australia, lacking the insect’s natural enemies, where more intense negative effects on pine species may occur. To date, Aleppo pine’s physiological responses to the infestation by M. hellenica are largely unknown. This study aimed at assessing the responses of Aleppo pine to the giant pine scale attack, both on an ecophysiological and a metabolic level. For this purpose, gas exchange, needle water status, and carbon and nitrogen content were measured during 1 year on healthy and infested adult trees. M etabolic profiling of Aleppo pine needles was also performed before, during, and after the high feeding activity of the insect. The maintenance of stable relative water content, δ13C signatures, and chlorophyll fluorescence in the needles of infested pines indicated that infestation did not induce drought stress to the host pines. At the peak of infestation, stomatal closure and a pronounced reduction in assimilation were observed and were associated with the accumulation of sugars in the needles, probably due to impaired phloem loading. At the end of the infestation period, tricarboxylic acids were induced and phenolic compounds were enhanced in the needles of infested pines. These metabolic responses, together with the recovery of photosynthesis after the end of M. hellenica intense feeding, indicate that in the studied region and under the current climate, Aleppo pine is resilient to the infestation by the giant pine scale. Future research should assess whether these promising defense mechanisms are also employed by other host pines, particularly in regions of the world recently invaded by the giant pine scale, as well as under more xerothermic regimes.

Plant growth strategies are adapted to resource availability in native habitat(s), and thus reflected in traits such as photosynthetic and respiratory capacities of leaves. We explored acclimation of such traits to contrasting light and nitrogen supply for Flooded Gum (Eucalyptus grandis) and Mountain Ash (E. regnans) saplings growing under warm-temperate conditions. We focused on respiration parameters that are not routinely measured, but could expand our mechanistic understanding of trait correlation networks [interdependencies between leaf mass per area (LMA), foliar nitrogen concentration (Nmass), photosynthetic capacity (mass- based Amax) and respiration (R)]. We measured temperature responses of leaf R via calorimetric methods, in order to derive three respiration parameters that are useful to characterize capacity and flux mode of respiratory oxygen reduction in mature source leaves and young sink leaves. Subtropical E. grandis saplings produced 3-fold more biomass than E. regnans, and responded more strongly to enhanced supplies of nitrogen, in particular under semi-shade conditions. Acclimation of LMA to growth irradiance was more plastic in E. grandis, but light treatment had no effect on Nmass and mass-based Amax in this species. In E. regnans, growth under abundant nitrogen and full sunlight caused significant increases in foliar Nmass–albeit not matched by relatively modest increases in mass-based Amax. Cool-temperate E. regnans saplings appeared to allocate a substantial share of leaf-N to protective functions upon exposure to full sunlight. Foliar nitrogen was used more effectively for the production of new foliage in E. grandis, owing to better coordination with foliar capacity and flux mode of mitochondrial oxygen reduction. Most of the variation in three respiration parameters can be explained by a few physiological/anatomical variables, responding to variation in absolute and relative demand for ATP, reducing power and anabolic intermediates. These demands strongly depended on leaf developmental stage and also varied between species and treatments. Our approach opens a path to improved process- based understanding of respiratory flux control, which could facilitate predictions of plant respiration in a changing environment.

The paradigm that permafrost-affected soils show restricted mineral nitrogen (N) cycling in favor of organic N compounds is based on the observation that net N mineralization rates in these cold climates are negligible. However, we find here that this perception is wrong. By synthesizing published data on N cycling in the plant-soil-microbe system of permafrost ecosystems we show that gross ammonification and nitrification rates in active layers were of similar magnitude and showed a similar dependence on soil organic carbon (C) and total N concentrations as observed in temperate and tropical systems. Moreover, high protein depolymerization rates and only marginal effects of C:N stoichiometry on gross N turnover provided little evidence for N limitation. Instead, the rather short period when soils are not frozen is the single main factor limiting N turnover. High gross rates of mineral N cycling are thus facilitated by released protection of organic matter in active layers with nitrification gaining particular importance in N-rich soils, such as organic soils without vegetation. Our finding that permafrost-affected soils show vigorous N cycling activity is confirmed by the rich functional microbial community which can be found both in active and permafrost layers. The high rates of N cycling and soil N availability are supported by biological N fixation, while atmospheric N deposition in the Arctic still is marginal except for fire-affected areas. In line with high soil mineral N production, recent plant physiological research indicates a higher importance of mineral plant N nutrition than previously thought. Our synthesis shows that mineral N production and turnover rates in active layers of permafrost-affected soils do not generally differ from those observed in temperate or tropical soils. We therefore suggest to adjust the permafrost N cycle paradigm, assigning a generally important role to mineral N cycling. This new paradigm suggests larger permafrost N climate feedbacks than assumed previously.

The present study elucidated whether roots of temperate forest trees can take up organic phosphorus in the form of ATP. Detached non-mycorrhizal roots of beech (Fagus sylvatica) and gray poplar (Populus x canescens) were exposed under controlled conditions to 33P-ATP and/or 13C/15N labeled ATP in the presence and absence of the acid phosphatase inhibitor MoO42-. Accumulation of the respective label in the roots was used to calculate 33P, 13C and 15N uptake rates in ATP equivalents for comparison reason. The present data shown that a significant part of ATP was cleaved outside the roots before phosphate (Pi) was taken up. Furthermore, nucleotide uptake seems more reasonable after cleavage of at least one Pi unit as ADP, AMP and/or as the nucleoside adenosine. Similar results were obtained when still attached mycorrhizal roots of adult beech trees and their natural regeneration of two forest stands were exposed to ATP in the presence or absence of MoO42-. Cleavage of Pi from ATP by enzymes commonly present in the rhizosphere, such as extracellular acid phosphatases, ecto-apyrase and/or nucleotidases, prior ADP/AMP/adenosine uptake is highly probable but depended on the soil type and the pH of the soil solution. Although uptake of ATP/ADP/AMP cannot be excluded, uptake of the nucleoside adenosine without breakdown into its constituents ribose and adenine is highly evident. Based on the 33P, 13C, and 15N uptake rates calculated as equivalents of ATP the ‘pro and contra’ for the uptake of nucleotides and nucleosides is discussed.Short SummaryRoots take up phosphorus from ATP as Pi after cleavage but might also take up ADP and/or AMP by yet unknown nucleotide transporter(s) because at least the nucleoside adenosine as N source is taken up without cleavage into its constituents ribose and adenine.

The aim of present study was to elucidate the significance of the phosphorylated pathway of Ser production for Cys biosynthesis in leaves at day and night and upon cadmium (Cd) exposure. For this purpose, Arabidopsis wildtype plants as control and its psp mutant knocked-down in phosphoserine phosphatase (PSP) were used to test if (i) photorespiratory Ser is the dominant precursor of Cys synthesis in autotrophic tissue in the light, (ii) the phosphorylated pathway of Ser production can take over Ser biosynthesis in leaves at night, and (iii) Cd exposure stimulates Cys and glutathione (GSH) biosynthesis and effects the crosstalk of S and N metabolism, irrespective of the Ser source. Glycine (Gly) and Ser contents were not affected by reduction of the psp transcript level confirming that the photorespiratory pathway is the main route of Ser synthesis. The reduction of the PSP transcript level in the mutant did not affect day/night regulation of sulfur fluxes while day/night fluctuation of sulfur metabolite amounts were no longer observed, presumably due to slower turnover of sulfur metabolites in the mutant. Enhanced contents of non-protein thiols in both genotypes and of GSH only in the psp mutant were observed upon Cd treatment. Mutation of the phosphorylated pathway of Ser biosynthesis caused an accumulation of alanine, aspartate, lysine and a decrease of branched-chain amino acids. Knock-down of the PSP gene induced additional defense mechanisms against Cd toxicity that differ from those of WT plants.

This study was aimed at elucidating the significance of photorespiratory serine (Ser) production for cysteine (Cys) biosynthesis. For this purpose, sulfur (S) metabolism and its crosstalk with nitrogen (N) and carbon (C) metabolism were analyzed in wildtype Arabidopsis and its photorespiratory bou-2 mutant with impaired glycine decarboxylase (GDC) activity. Foliar glycine and Ser contents were enhanced in the mutant at day and night. The high Ser levels in the mutant cannot be explained by transcript abundances of genes of the photorespiratory pathway or two alternative pathways of Ser biosynthesis. Despite enhanced foliar Ser, reduced GDC activity mediated a decline in sulfur flux into major sulfur pools in the mutant, as a result of deregulation of genes of sulfur reduction and assimilation. Still, foliar Cys and glutathione contents in the mutant were enhanced. The use of Cys for methionine and glucosinolates synthesis was reduced in the mutant. Reduced GDC activity in the mutant downregulated Calvin Cycle and nitrogen assimilation genes, upregulated key enzymes of glycolysis and the tricarboxylic acid (TCA) pathway and modified accumulation of sugars and TCA intermediates. Thus, photorespiratory Ser production can be replaced by other metabolic Ser sources, but this replacement deregulates the cross-talk between S, N, and C metabolism.

The temperate climax tree species Fagus sylvatica and the floodplain tree species Populus × canescens possess contrasting phosphorus (P) nutrition strategies. While F. sylvatica has been documented to display P storage and mobilization (Netzer et al., 2017), this was not observed for Populus × canescens (Netzer et al., 2018b). Nevertheless, changes in the abundance of organic bound P in gray poplar trees indicated adaptation of the P nutrition to different needs during annual growth. The present study aimed at characterizing seasonal changes in metabolite and lipid abundances in gray poplar and uncovering differences in metabolite requirement due to specific needs depending on the season. Seasonal variations in the abundance of (i) sugar-Ps and phospholipids, (ii) amino acids, (iii) sulfur compounds, and (iv) carbon metabolites were expected. It was hypothesized that seasonal changes in metabolite levels relate to N, S, and C storage and mobilization. Changes in organic metabolites binding Pi (Porg) are supposed to support these processes. Variation in triacylglycerols, in sugar-phosphates, in metabolites of the TCA cycle and in the amino acid abundance of poplar twig buds, leaves, bark, and wood were found to be linked to changes in metabolite abundances as well as to C, N, and S storage and mobilization processes. The observed changes support the view of a lack of any P storage in poplar. Yet, during dormancy, contents of phospholipids in twig bark and wood were highest probably due to frost-hardening and to its function in extra-plastidic membranes such as amyloplasts, oleosomes, and protein bodies. Consistent with this assumption, in spring sugar-Ps increased when phospholipids declined and poplar plants entering the vegetative growth period and, hence, metabolic activity increases. These results indicate that poplar trees adopt a policy of P nutrition without P storage and mobilization that is different from their N- and S-nutrition strategies.

Phosphorus (P) is one of the most important macronutrients limiting plant growth and development, particularly in forest ecosystems such as temperate beech (Fagus sylvatica) forests in Central Europe. Efficient tree internal P cycling during annual growth is an important strategy of beech trees to adapt to low soil-P. Organic P (Porg) is thought to play a decisive role in P cycling, but the significance of individual compounds and processes has not been elucidated. To identify processes and metabolites involved in P cycling of beech trees, polar-metabolome and lipidome profiling was performed during annual growth with twig tissues from a sufficient (Conventwald, Con) and a low-soil-P (Tuttlingen, Tut) forest. Autumnal phospholipid degradation in leaves and P export from senescent leaves, accumulation of phospholipids and glucosamine-6-phosphate (GlcN6P) in the bark, storage of N-acetyl-D-glucosamine-6-phosphate (GlcNAc6P) in the wood, and establishing of a phospholipid “start-up capital” in buds constitute main processes involved in P cycling that were enhanced in beech trees on low-P soil of the Tut forest. In spring, mobilization of P from storage pools in the bark contributed to an effective P cycling. Due to the higher phospholipid “start-up capital” in buds of Tut beeches, the P metabolite profile in developing leaves in spring was similar in beech trees of both forests. During summer, leaves of Tut beeches meet their phosphate (Pi) needs by replacing phospholipids by galacto- and sulfolipids. Thus, several processes contribute to adequate Pi supply on P impoverished soil thereby mediating similar growth of beech at low and sufficient soil-P availability.

Different plant species were investigated from two Aeolian Islands located in close vicinity, one with fumarolic activity (Vulcano) and one without (Lipari). On Vulcano, elevated concentrations of SO2/H2S determined in ambient air indicated the need of plants to adapt to harmful sulphur concentrations by detoxification strategies. The current study was focused on evaluating the element composition of plant leaves in relation to soil mineral contents. The soil of Volcano was characterised by a significantly lower pH on all three sampling sites as well as very high amounts of sulphur and plant available sulphate due to volcanic activities, compared to Lipari. By contrast, a general difference in the composition of trace elements in the soil was not observed between the islands, apart from arsenic, which was increased at all three sampling sites on Vulcano. Element accumulation in the leaves differed between the two islands. The tested species showed a significant higher accumulation of numerous elements (Al, B, Fe, K, Mg, Mn, Ni, and Zn) on Vulcano compared to Lipari, while excluding Ca and Mo. These differences in element accumulation in the leaves between the islands may be caused by the lower soil pH on Vulcano. Extreme sulphur accumulation was found for all tested species on Vulcano, but was lower in woody species with higher dry matter content compared to herbaceous species with lower dry matter content. This caused a significantly negative correlation between plant sulphur and dry matter content. From these results, it is concluded that species with higher dry matter contents possess a more effective protection against extreme sulphur accumulation. Strategies to cope with other potentially toxic elements in the soil ranged from exclusion to hyper-accumulation. Hierarchical cluster analyses of the leaf element content revealed a clear separation between two groups: First, herbaceous perennial plants as strong accumulators; and second, woody perennial plants such as shrubs or trees as less strong accumulators, with the primordial species Fumaria capreolata representing an outside group.

Replacement of phospholipids with phosphorus (P)-free lipids in cellular membranes has been identified as a mechanism facilitating fast rates of photosynthesis when phosphorus availability is limited. We measured photosynthetic rates, leaf and soil P fractions, and foliar membrane lipid compositions for five species (Geranium antrorsum, Ranunculus graniticola, Poa costiniana, Poa hiemata, and Veronica derwentiana) common to two Australian subalpine ecosystems of contrasting parent material to characterize the extent to which they have adapted to long-term P availability. Our results indicate limited tolerance to reduced P, albeit adaptation strategies differ among species. Under reduced P conditions, phospholipids were replaced in foliage by galactolipids and sulfolipids, but photosynthesis was still impaired owing to reduced stomatal conductance. Accumulation of antioxidants, including carotenoids and alpha-tocopherol, in leaves with limited P supply suggests oxidative stress. Our field study shows that while subalpine Australian plants of a variety of life forms adapt to P availability by replacing phospholipids with P-free lipids in foliar membranes, this adaptation is insufficient to fully mitigate the effects of reduced P on photosynthesis.

The coniferous forest tree Douglas-fir (Pseudotsuga menziesii) is native to the pacific North America, and is increasingly planted in temperate regions worldwide. Nitrogen (N) metabolism is of great importance for growth, resistance and resilience of trees. In the present study, foliar N metabolism of adult trees of three coastal and one interior provenance of Douglas-fir grown at two common gardens in southwestern Germany (Wiesloch, W; Schluchsee, S) were characterized in two subsequent years. Both the native North American habitats of the seed sources and the common garden sites in Germany differ in climate conditions. Total and mineral soil N as well as soil water content were higher in S compared to W. We hypothesized that i) provenances differ constitutively in N pool sizes and composition, ii) N pools are affected by environmental conditions, and iii) that effects of environmental factors on N pools differ among interior and coastal provenances. Soil water content strongly affected the concentrations of total N, soluble protein, total amino acids (TAA), arginine and glutamate. Foliar concentrations of total N, soluble protein, structural N and TAA of trees grown at W were much higher than in trees at S. Provenance effects were small but significant for total N and soluble protein content (interior provenance showed lowest concentrations), as well as arginine, asparagine and glutamate. Our data suggest that needle N status of adult Douglas-fir is independent from soil N availability and that low soil water availability induces a re-allocation of N from structural N to metabolic N pools. Small provenance effects on N pools suggest that local adaptation of Douglas-fir is not dominated by N conditions at the native habitats.

Biotic stress by mass outbreaks of defoliating pest insects does not only affect tree performance by reducing its photosynthetic capacity, but also changes N cycling in the soil of forest ecosystems. However, how insect induced defoliation affects soil N fluxes and, in turn, tree N nutrition is not well-studied. In the present study, we quantified N input and output fluxes via dry matter input, throughfall, and soil leachates. Furthermore, we investigated the effects of mass insect herbivory on tree N acquisition (i.e., organic and inorganic 15N net uptake capacity of fine roots) as well as N pools in fine roots and needles in a Scots pine (Pinus sylvestris L.) forest over an entire vegetation period. Plots were either infested by the nun moth (Lymantria monacha L.) or served as controls. Our results show an increased N input by insect feces, litter, and throughfall at the infested plots compared to controls, as well as increased leaching of nitrate. However, the additional N input into the soil did not increase, but reduce inorganic and organic net N uptake capacity of Scots pine roots. N pools in the fine roots and needles of infested trees showed an accumulation of total N, amino acid-N, protein-N, and structural N in the roots and the remaining needles as a compensatory response triggered by defoliation. Thus, although soil N availability was increased via surplus N input, trees did not respond with an increased N acquisition, but rather invested resources into defense by accumulation of amino acid-N and protein-N as a survival strategy.

Plants adapt to the environment by either long-term genome evolution or by acclimatization processes where the cellular processes and metabolism of the plant are adjusted within the existing potential in the genome. Here we studied the adaptation strategies in date palm, Phoenix dactylifera, under mild heat, drought and combined heat and drought by transcriptomic and metabolomic profiling. In transcriptomics data, combined heat and drought resembled heat response, whereas in metabolomics data it was more similar to drought. In both conditions, soluble carbohydrates, such as fucose, and glucose derivatives, were increased, suggesting a switch to carbohydrate metabolism and cell wall biogenesis. This result is consistent with the evidence from transcriptomics and cis-motif analysis. In addition, transcriptomics data showed transcriptional activation of genes related to reactive oxygen species in all three conditions (drought, heat, and combined heat and drought), suggesting increased activity of enzymatic antioxidant systems in cytosol, chloroplast and peroxisome. Finally, the genes that were differentially expressed in heat and combined heat and drought stresses were significantly enriched for circadian and diurnal rhythm motifs, suggesting new stress avoidance strategies.

European beech forests growing on marginal calcareous soils have been proposed to be vulnerable to decreased soil water availability. This could result in a large-scale loss of ecological services and economical value in a changing climate. In order to evaluate the potential consequences of this drought-sensitivity, we investigated potential species range shifts for European beech forests on calcareous soil in the 21st century by statistical species range distribution modelling for present day and projected future climate conditions. We found a dramatic decline by 78% until 2080. Still the physiological or biogeochemical mechanisms underlying the drought sensitivity of European beech are largely unknown. Drought sensitivity of beech is commonly attributed to plant physiological constraints. Furthermore, it has also been proposed that reduced soil water availability could promote nitrogen (N) limitation of European beech due to impaired microbial N cycling in soil, but this hypothesis has not yet been tested. Hence we investigated the influence of simulated climate change (increased temperatures, reduced soil water availability) on soil gross microbial N turnover and plant N uptake in the beech-soil interface of a typical mountainous beech forest stocking on calcareous soil in SW Germany. For this purpose, triple 15N isotope labelling of intact beech seedling-soil-microbe systems was combined with a space-for-time climate change experiment. We found that nitrate was the dominant N source for beech natural regeneration. Reduced soil water content caused a persistent decline of ammonia oxidizing bacteria and therefore, a massive attenuation of gross nitrification rates and nitrate availability in the soil. Consequently, nitrate and total N uptake of beech seedlings were strongly reduced so that impaired growth of beech seedlings was observed already after one year of exposure to simulated climatic change. We conclude that the N cycle in this ecosystem and here specifically nitrification is vulnerable to reduced water availability, which can directly lead to nutritional limitations of beech seedlings. This tight link between reduced water availability, drought stress for nitrifiers, decreased gross nitrification rates and nitrate availability and finally nitrate uptake by beech seedlings could represent the Achilles' heel for beech under climate change stresses.

The performance of European beech (Fagus sylvatica L.) is severely threatened by abiotic and biotic stresses, but the resilience of its provenances from distinct geographic areas has not been sufficiently studied. Therefore, the performance of European beech was investigated in an international provenance trial consisting of twenty-one natural populations originating from Central Europe and Southeast Europe located on Medvednica Mountain (Croatia). The performance of European beech was investigated by characterizing I) damage types, II) crown damage intensity, III) damage frequency, and IV) clustering of provenances based on damage types. Anthracnose, galls, chewing damages, and aphids were recorded on leaves, and canker on the trunk and branches. The crown damage intensity was minute (less than 10%) for all types of disease. Anthracnose was the most common damage, followed in descending order by galls, canker, chewing damages and aphids. When the types of damage were analysed using descriptive statistics (frequency analysis and clustering), significant differences were recorded between provenances (χ2=322.19, p

The present study with young poplar trees aimed at characterizing the effect of O2 shortage in the soil on net uptake of NO3- and NH4+ and the spatial distribution of the N taken up. Moreover, we assessed biomass increment as well as N status of the trees affected by O2 deficiency. For this purpose, an experiment was conducted in which hydroponically grown young poplar trees were exposed to hypoxic and normoxic (control) conditions for 14 days. 15N-labelled NO3- and NH4+ were used to elucidate N uptake and distribution of currently absorbed N and N allocation rates in the plants. Whereas shoot biomass was not affected by soil O2 deficiency, it significantly reduced root biomass and, consequently, the root-to-shoot ratio. Uptake of NO3- but not of NH4+ by the roots of the trees was severely impaired by hypoxia. As a consequence of reduced N uptake, the N content of all poplar tissues was significantly diminished. Under normoxic control conditions, the spatial distribution of currently absorbed N and N allocation rates differed depending on the N source. Whereas NO3- derived N was mainly transported to the younger parts of the shoot, particularly to the developing and young mature leaves, N derived from NH4+ was preferentially allocated to older parts of the shoot, mainly to wood and bark. Soil O2 deficiency enhanced this differential allocation pattern. From these results we assume that NO3- was assimilated in developing tissues and preferentially used to maintain growth and ensure plant survival under hypoxia, whereas NH4+ based N was used for biosynthesis of storage proteins in bark and wood of the trees. Still, further studies are needed to understand the mechanistic basis as well as the eco-physiological advantages of such differential allocation patterns.

Climate change poses direct or indirect influences on physiological mechanisms in plants. In particular, long living plants like trees have to cope with the predicted climate changes (i.e. drought and air warming) during their life span. The present study aimed to quantify the consequences of simulated climate change for foliar N metabolites over a drought-rewetting-drought course. Saplings of three Central European oak species (i.e. Quercus robur, Q. petraea, Q. pubescens) were tested on two different soil types (i.e. acidic and calcareous). Consecutive drought periods increased foliar amino acid-N and soluble protein-N concentrations at the expense of structural N in all three oak species. In addition, transient effects on foliar metabolite dynamics were observed over the drought-rewetting-drought course. The lowest levels of foliar soluble protein-N, amino acid-N and potassium cation with a minor response to drought and air warming were found in the oak species originating from the driest/warmest habitat (Q. pubescens) compared to Q. robur and Q. petraea. Higher foliar osmolyte-N and potassium under drought and air warming were observed in all oak species when grown on calcareous versus acidic soil. These results indicate that species-specific differences in physiological mechanisms to compensate drought and elevated temperature are modified by soil acidity.

In the future, periods of strongly increased temperature in concert with drought (heat waves) will have potentially detrimental effects on trees and forests in Central Europe. Norway spruce might be at risk in the future climate of Central Europe. However, Douglas-fir is often discussed as an alternative for the drought and heat sensitive Norway spruce, because some provenances are considered to be well adapted to drier and warmer conditions. In this study, we identified the physiological and growth responses of seedlings from two different Douglas-fir provenances to increased temperature and atmospheric drought during a period of 92 days. We analysed (i) plant biomass, (ii) carbon stable isotope composition as an indicator for time integrated intrinsic water use efficiency, (iii) apparent respiratory carbon isotope fractionation as well as (iv) the profile of polar low molecular metabolites. Plant biomass was only slightly affected by increased temperatures and atmospheric drought but the more negative apparent respiratory fractionation indicated a temperature-dependent decrease in the commitment of substrate to the tricarboxylic acid cycle. The metabolite profile revealed that the simulated heat wave induced a switch in stress protecting compounds from proline to polyols. We conclude that metabolic acclimation successfully contributes to maintain functioning and physiological activity in seedlings of both Douglas-fir provenances under conditions that are expected during heat waves (i.e. elevated temperatures and atmospheric drought). Douglas-fir might be a potentially important tree species for forestry in Central Europe under changing climatic conditions.

Modern agriculture is based on the notion that nitrate is the main source of nitrogen (N) for crops, but nitrate is also the most mobile form of N and easily lost from soil. Efficient acquisition of nitrate by crops is therefore a prerequisite for avoiding off-site N pollution. Sugarcane is considered the most suitable tropical crop for biofuel production, but surprisingly high N fertilizer applications in main producer countries raise doubt about the sustainability of production and are at odds with a carbon-based crop. Examining reasons for the inefficient use of N fertilizer, we hypothesized that sugarcane resembles other giant tropical grasses which inhibit the production of nitrate in soil and differ from related grain crops with a confirmed ability to use nitrate. The results of our study support the hypothesis that N-replete sugarcane and ancestral species in the Andropogoneae supertribe strongly prefer ammonium over nitrate. Sugarcane differs from grain crops, sorghum and maize, which acquired both N sources equally well, while giant grass, Erianthus, displayed an intermediate ability to use nitrate. We conclude that discrimination against nitrate and a low capacity to store nitrate in shoots prevents commercial sugarcane varieties from taking advantage of the high nitrate concentrations in fertilized soils in the first three months of the growing season, leaving nitrate vulnerable to loss. Our study addresses a major caveat of sugarcane production and affords a strong basis for improvement through breeding cultivars with enhanced capacity to use nitrate as well as through agronomic measures that reduce nitrification in soil.

ABSTRACT Previous ecological studies of dioecious plant species have found that female plants preferentially occur at lower altitudes where there are typically better nutritional conditions, while male plants often occur in less favorable sites. We compared the ecophysiological performance of male and female plants in three populations of the dioecious Baccharis concinna, an endemic species of rupestrian grasslands of Serra do Cipó, in southeastern Brazil. We hypothesized that physiological differences between the sexes would explain the distribution patterns of the populations. Analyses of the tissue content of phosphorus (P), calcium (Ca), potassium (K) and sodium (Na), and carbon and nitrogen stable isotopes, were used to assess nutritional status and water use efficiency (WUE) in plant leaves, stems and roots of male and female plants in three populations located along an elevational gradient. Differences among populations were related to decreased nutrient levels and WUE at higher elevations, but an effect of sex was found only for %C, with male plants having slightly higher values. In conclusion, the sex ratios in the studied populations of B. concinna could not be attributed to differences in nutrient acquisition and WUE.