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Ricinus communis is an important oilseed crop worldwide and is also considered one of the best potential plants for salt‐affected soil improvement in northeast China. However, little is known about photosynthesis and carbohydrate metabolism in this plant, nor the distribution of carbohydrates in cotyledons and roots under salinity stress.In the present study, seedling growth, gas exchange parameters (PN, E, gs and Ci), carbohydrate (fructose, sucrose, glucose, soluble sugar and starch) metabolism and related enzymes and genes were measured in Ricinus plants.Under salt stress, PN of cotyledons decreased significantly (P < 0.05), resulting in weak photosynthetic capacity. Furthermore, salt stress increased sucrose and glucose content in cotyledons, but decreased soluble sugar and starch content. However, sucrose increased and starch decreased in roots. This may be correlated with the increasing sugar metabolism under salinity, including notable changes in sugar‐related enzyme activities (SPS, SuSy, α‐amylase and β‐amylase) and gene expression of RcINV, RcSUS, RcAmY, RcBAM and RcGBE1.The results suggest that salinity reduces photosynthesis of cotyledons, alters carbohydrate allocation between cotyledons and roots and also promotes starch utilization in cotyledons and starch biosynthesis in roots, leading to a functional imbalance between cotyledons and roots. Together, these findings provide insights into the crucial role of sugar metabolism in improving salt‐tolerance of Ricinus during the early seedling growth stage. [ABSTRACT FROM AUTHOR]

Understanding how plants adjust their requirements for different N forms can help elucidate plant coexistence strategies in N‐limited desert ecosystems. To understand the mechanisms involved, we investigated whether two desert herbs can directly absorb dissolved organic nitrogen (N) and tested whether the patterns changed over different growth stages.Two dominant herbaceous species, Astragalus arpilobus and Arnebia decumbens, from the southern edge of the Gurbantunggut desert, China, were selected. Short‐term (24 h) 15N‐labelled tracer (15N‐NO3, 15N‐NH4, 2‐13C‐15N‐Glycine) treatments were conducted at two soil depths (0–5 cm and 5–15 cm) in the season of rapid growth (June) and in the peak biomass season (July). Enrichment in 13C and 15N was assessed in the two species receiving glycine.The ratio 13C:15N was 0.21–1.39 at the 24‐h harvest, suggesting that approximately 10.5–69.5% of glycine had been absorbed. The amount of absorbed 15N was significantly affected by species, month, soil depth and N form. The two species absorbed most 15N from the 0–5 cm soil layer, and the absorption rate in July was higher than that in June. The absorption of 15N‐NO3 and 15N‐NH4 was significantly higher than that of 2‐13C‐15N‐Glycine.The results indicate that these herbs could use amino acids in the N‐deficient desert ecosystem. The two co‐existing species used different forms of inorganic N for their requirements and maintained a specific preference throughout various growth stages. [ABSTRACT FROM AUTHOR]

Growth and functioning of Sphagnum mosses are closely linked to water level and chemistry. Sphagnum mosses occur in wet, generally acidic conditions, and when buffered, alkaline water is known to negatively impact Sphagnum. The effects of time, dose and species‐specific responses of buffered, alkaline water on Sphagnum are largely unknown.We investigated the effects of bicarbonate and calcium on the survival, growth and physiological functioning of seven Sphagnum species occurring in contrasting environments, from raised bogs to (rich) fens. Mosses were submerged in different concentrations of bicarbonate and calcium solutions for 10 weeks under climate‐controlled circumstances.After 2 weeks, all species exposed to the high bicarbonate treatment (2.0 mM) showed severe potassium leakage and swift discoloration. In contrast, species showed differential responses to the intermediate bicarbonate treatment (0.8 mM), some with a later onset of potassium leakage. S. squarrosum, S. teres & S. contortum generally persisted the longest, with all species dying after 6 to 10 weeks. Calcium alone, in contrast, negatively affected S. squarrosum, S. teres & S. contortum, causing discoloration and potassium leakage.Our study shows enrichment with bicarbonate, but not calcium, is detrimental for most Sphagnum species tested. A mechanistic model was developed that is consistent with dose and duration dependence and the species specificity. Future conservation and restoration measures for Sphagnum‐dominated habitats and Sphagnum farming (cultivation, production and harvest of Sphagnum moss biomass) should limit flooding with bicarbonate‐rich waters while investigating new management options, like acidifying surface waters to lower bicarbonate levels. [ABSTRACT FROM AUTHOR]

Persistence and colonization by tree species in an environment following a fire depends on the effects on seed germination and seedling development. We used seeds of Kielmeyera coriacea and Qualea parviflora as a model to test the effects of high temperatures on germination and initial development of tree seedlings.We exposed the seeds to heat flow (70, 100, 130, 150 or 170 °C) for 2 or 5 min and compared the germination with that of unheated seeds (control). Seedlings were then harvested after 3, 7 or 15 days to evaluate aerial and root mass, root:shoot ratio, presence of cotyledon opening, true leaves, and secondary roots.We found no effect on germination for seeds exposed to temperatures ≥150 °C. However, germination was significantly reduced for seeds exposed to 100 °C for both 2 and 5 min. The mass of 15‐day‐old K. coriacea seedlings was smaller when seeds were heated at 70 °C for 5 min or at temperatures higher or equal to 100 °C. Qualea parviflora seedlings did not show any difference in mass, but there were marginal differences in the presence of roots and the opening of cotyledons. Kielmeyera coriacea seedlings allocated biomass faster than Q. parviflora.High temperatures affect both quantity and quality of germinable seeds, as well as biomass allocation during initial seedling development. These factors may explain the decrease in seedlings observed after fire, suggesting a bottleneck effect that influences population dynamics and species persistence in systems with frequent fires. [ABSTRACT FROM AUTHOR]

Biochar (BC) amendment has been proposed as a promising strategy for mitigating greenhouse gas (GHG) emissions, specifically carbon dioxide (CO 2 ), methane (CH 4 ), and nitrous oxide (N 2 O). Conducting a meta-analysis to evaluate the impact of biochar on microbial genetic profile, community structure, and phospholipid fatty acid (PLFA) contents can aid in identifying key microbial groups involved in GHG production and consumption, and assessing the overall effectiveness of biochar in reducing GHG emissions. The present meta-analysis revealed that the addition of biochar resulted in a 22 % and 41 % reduction in pmoA and mcrA genes of methanogenic microorganisms, respectively. The mcrA/pmoA ratio significantly increased by 81 %. Gene abundances exhibited a positive response to biochar amendment, with increases observed in nifH, nirK, nirS, nosZ, and nosZ (nirS + nirK) genes by 13 %, 32 %, 37 %, 42 %, and 79 %, respectively. Moreover, biochar amendment influenced the microbial community structure accordingly. The concentration of PLFAs increased in response to BC treatment in the following order: A-bacteria (+49 %) < Fungi (+30 %) < Gram-pb (+21 %) < G-bacteria (+17 %) < Gram-nb (+11 %). These findings indicate that biochar amendment shapes the microbial community structure, further emphasizing its significance in enhancing soil fertility., (Copyright © 2024. Published by Elsevier B.V.)

Both sulfur (S) supply and legume-rhizobium symbiosis can significantly contribute to enhancing the efficiency of phytoremediation of heavy metals (HMs). However, the regulatory mechanism determining the performance of legumes at lead (Pb) exposure have not been elucidated. Here, we cultivated black locust (Robinia pseudoacacia L.), a leguminous woody pioneer species at three S supply levels (i.e., deficient, moderate, and high S) with rhizobia inoculation and investigated the interaction of these treatments upon Pb exposure. Our results revealed that the root system of Robinia has a strong Pb accumulation and anti-oxidative capacity that protect the leaves from Pb toxicity. Compared with moderate S supply, high S supply significantly increased Pb accumulation in roots by promoting the synthesis of reduced S compounds (i.e., thiols, phytochelatin), and also strengthened the antioxidant system in leaves. Weakened defense at deficient S supply was indicated by enhanced oxidative damage. Rhizobia inoculation alleviated the oxidative damage of its Robinia host by immobilizing Pb to reduce its absorption by root cells. Together with enhanced Pb chelation in leaves, these mechanisms strengthen Pb detoxification in the Robinia-rhizobia symbiosis. Our results indicate that appropriate S supply can improve the defense of legume-rhizobia symbiosis against HM toxicity., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Increasing desertification constitutes a global environmental problem, mainly driven by climate change and inappropriate land-use that limits agriculture, forestry and human colonization. The selection of suitable plant species to mitigate desertification is particularly challenging, as it usually requires simultaneous counteraction against a whole set of unfavourable environmental conditions, including heat, drought, high tropospheric ozone and salinity. It therefore seems useful to identify the survival strategies of plants native in desert environments. Date palm constitutes a plant species native in desert environments and cultivated worldwide in arid regions that have been studied intensively for stress defence during the last decade. The present review summarizes the current state of biochemical stress defence mechanisms including avoidance, osmotic and metabolic adjustments and reactive oxygen species scavenging, addresses whole-plant regulations and trade-off between stress compensation/defence and growth of date palms. The review advances our knowledge about how this typical desert species copes with both individual and multiple environmental stresses at the cellular to the whole-plant level, and identifies areas of future research required to fully understand the strategies of this plant species to survive in the desert, thereby contributing to efforts for the mitigation of climate change and desertification., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

In their environment, plants are exposed to a multitude of abiotic and biotic stresses that differ in intensity, duration and severity. As sessile organisms, they cannot escape these stresses, but instead have developed strategies to overcome them or to compensate for the consequences of stress exposure. Defence can take place at different levels and the mechanisms involved are thought to differ in efficiency across these levels. To minimise metabolic constraints and to reduce the costs of stress defence, plants prioritise first-line defence strategies in the apoplastic space, involving ascorbate, defensins and small peptides, as well as secondary metabolites, before cellular processes are affected. In addition, a large number of different symplastic mechanisms also provide efficient stress defence, including chemical antioxidants, antioxidative enzymes, secondary metabolites, defensins and other peptides as well as proteins. At both the symplastic and the apoplastic level of stress defence and compensation, a number of specialised transporters are thought to be involved in exchange across membranes that still have not been identified, and information on the regeneration of different defence compounds remains ambiguous. In addition, strategies to overcome and compensate for stress exposure operate not only at the cellular, but also at the organ and whole-plant levels, including stomatal regulation, and hypersensitive and systemic responses to prevent or reduce the spread of stress impacts within the plant. Defence can also take place at the ecosystem level by root exudation of signalling molecules and the emission of volatile organic compounds, either directly or indirectly into the rhizosphere and/or the aboveground atmosphere. The mechanisms by which plants control the production of these compounds and that mediate perception of stressful conditions are still not fully understood. Here we summarise plant defence strategies from the cellular to ecosystem level, discuss their advantages and disadvantages for plant growth and development, elucidate the current state of research on the transport and regeneration capacity of defence metabolites, and outline insufficiently explored questions for further investigation., (© 2024 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Lavender plantation is globally expanded due to the increasing demand of its essential oil and its popularity as an ornamental species. However, lavender plantations, and consequently essential oil industries, are threatened by more frequent and severe drought episodes in a globally changing climate. Still little is known about the changes in the general metabolome, which provides the precursors of essential oil production, by extended drought events. Prolonged drought fundamentally results in yield losses and changing essential oil composition. In the present study, the general metabolome of a main cultivated lavender species (Lavandula angustifolia Mill.) in response to water deprivation (WD) and re-watering was analyzed to identify the metabolomics responses. We found prolonged WD resulted in significant accumulations of glucose, 1,6-anhydro-β-D-glucose, sucrose, melezitose and raffinose, but declines of allulose, β-D-allose, altrose, fructose and D-cellobiose accompanied by decreased organic acids abundances. Amino acids and aromatic compounds of p-coumaric acid, hydrocaffeic acid and caffeic acid significantly accumulated at prolonged WD, whereas aromatics of cis-ferulic acid, taxifolin and two fatty acids (i.e., palmitic acid and stearic acid) significantly decreased. Prolonged WD also resulted in decreased abundances of polyols, particularly myo-inositol, galactinol and arabitol. The altered metabolite profiles by prolonged WD were mostly not recovered after re-watering, except for branched-chain amino acids, proline, serine and threonine. Our study illustrates the complex changes of leaf primary and secondary metabolic processes of L. angustifolia by drought events and highlights the potential impact of these precursors of essential oil production on the lavender industry., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Nitrogen (N 2 )-fixing legumes can be used for phytoremediation of toxic heavy metal Mercury (Hg) contaminated soil, but N 2 -fixation highly relies on phosphorus (P) availability for nodule formation and functioning. Here, we characterized the significance of P deficiency for Hg accumulation and toxicity in woody legume plants. Consequences for foliar and root traits of rhizobia inoculation, Hg exposure (+Hg) and low P (-P) supply, individually and in combination were characterized at both the metabolite and transcriptome levels in seedlings of two Robinia pseudoacacia L. provenances originating from contrasting climate and soil backgrounds, i.e., GS in northwest and the DB in northeast China. Our results reveal that depleted P mitigates the toxicity of Hg at the transcriptional level. In leaves of Robinia depleted P reduced oxidative stress and improved the utilization strategy of C, N and P nutrition; in roots depleted P regulated the expression of genes scavenging oxidative stress and promoting cell membrane synthesis. Rhizobia inoculation significantly improved the performance of both Robinia provenances under individual and combined +Hg and -P by promoting photosynthesis, increasing foliar N and P content and reducing H 2 O 2 and MDA accumulation despite enhanced Hg uptake. DB plants developed more nodules, had higher biomass and accumulated higher Hg amounts than GS plants and thus are suggested as the high potential Robinia provenance for future phytoremediation of Hg contaminated soils with P deficiency., Competing Interests: Declaration of Competing Interest The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Elsevier B.V. All rights reserved.)

The role of rhizobia in alleviating cadmium (Cd) stress in woody legumes is still unclear. Therefore, two types of black locust (Robinia pseudoacacia L.) with high and low Cd accumulation abilities were selected from 11 genotypes in China, and the effects of rhizobium (Mesorhizobium huakuii GP1T11) inoculation on the growth, CO 2 and H 2 O gas exchange parameters, Cd accumulation, and the absorption of mineral elements of the high (SX) and low Cd-accumulator (HB) were compared. The results showed that rhizobium-inoculation significantly increased biomass, shoot Cd contents, Cd accumulation, root-to-shoot translocation factor (TF) and the absorption and accumulation of mineral elements in both SX and HB. Rhizobium-inoculation increased chlorophyll a and carotenoid contents, and the intercellular carbon dioxide concentrations in HB plants. Under Cd exposure, the high-accumulator SX exhibited a significant decrease in photosynthetic CO 2 fixation (Pn) and an enhanced accumulation of Cd in leaves, but coped with Cd exposure by increasing chlorophyll synthesis, regulating stomatal aperture (Gs), controlling transpiration (Tr), and increasing the absorption and accumulation of mineral elements. In contrast, the low-accumulator HB was more sensitive to Cd exposure despite preferential accumulation of Cd in roots, with decreased chlorophyll and carotenoid contents, but significantly increased root biomass. Compared to the low-accumulator HB, non-inoculated Cd-exposed SX plants had higher chlorophyll contents, and rhizobium-inoculated Cd-exposed SX plants had higher Pn, Tr, and Gs as well as higher levels of P, K, Fe, Ca, Zn, and Cu. In conclusion, the high- and low-Cd-accumulator exhibited different physiological responses to Cd exposure. Overall, rhizobium-inoculation of black locust promoted the growth and heavy metal absorption, providing an effective strategy for the phytoremediation of heavy metal-contaminated soils by this woody legume., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Biochar could reshape microbial communities, thereby altering methylmercury (MeHg) concentrations in rice rhizosphere and seeds. However, it remains unclear whether and how biochar amendment perturbs microbe-mediated MeHg production in mercury (Hg) contaminated paddy soil. Here, we used pinecone-derived biochar and its six modified biochars to reveal the disturbance. Results showed that selenium- and chitosan-modified biochar significantly reduced MeHg concentrations in the rhizosphere by 85.83% and 63.90%, thereby decreasing MeHg contents in seeds by 86.37% and 75.50%. The two modified bicohars increased the abundance of putative Hg-resistant microorganisms Bacillus, the dominant microbe in rhizosphere. These reductions about MeHg could be facilitated by biochar sensitive microbes such as Oxalobacteraceae and Subgroup&#95;7. Pinecone-derived biochar increased MeHg concentration in rhizosphere but unimpacted MeHg content in seeds was observed. This biochar decreased the abundance in Bacillus but enhanced in putative Hg methylator Desulfovibrio. The increasing MeHg concentration in rhizosphere could be improved by biochar sensitive microbes such as Saccharimonadales and Clostridia. Network analysis showed that Saccharimonadales and Clostridia were the most prominent keystone taxa in rhizosphere, and the three biochars manipulated abundances of the microbes related to MeHg production in rhizosphere by those biochar sensitive microbes. Therefore, selenium- and chitosan-modified biochar could reduce soil MeHg production by these microorganisms, and is helpful in controlling MeHg contamination in rice., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Rhizobia and arbuscular mycorrhizal fungi (AMF) are symbiotic microorganisms important for plants grown in nutrient-deficient and heavy metal-contaminated soils. However, it remains unclear how plants respond to the coupled stress by heavy metal and nitrogen (N) deficiency under co-inoculation. Here, we investigated the synergistic effect of Mesorhizobium huakuii QD9 and Funneliformis mosseae on the response of black locust (Robinia pseudoacacia L.) grown in sand culture to cadmium (Cd) under N deficiency conditions. The results showed that single inoculation of AMF improved the growth and Cd resistance of black locust, co-inoculation improved the most. Compared to non-inoculated controls, co-inoculation mediated higher biomass and antioxidant enzyme activity, reduced oxidative stress, and promoted nodulation, mycorrhizal colonization, photosynthetic capacity, and N, P, Fe and Mg acquisition when exposed to Cd. This increase was significantly higher under N deficiency compared to N sufficiency. In addition, the uptake of Cd by co-inoculated black locust roots increased, but Cd translocation to the above-ground decreased under both N deficiency and sufficiency. Thus, in the tripartite symbiotic system, not merely metabolic processes but also Cd uptake increased under N deficiency. However, enhanced Cd detoxification in the roots and reduced allocation to the shoot likely prevent Cd toxicity and rather stimulated growth under these conditions., (© 2024 Scandinavian Plant Physiology Society.)

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., (© 2023 The Authors. The Plant Genome published by Wiley Periodicals LLC on behalf of Crop Science Society of America.)

The growth of fruit trees depends on the nitrogen (N) remobilization in mature tissues and N acquisition from the soil. However, in evergreen mature citrus (Citrus reticulata Blanco) leaves, proteins with N storage functions and hub molecules involved in driving N remobilization remain largely unknown. Here, we combined proteome and physiological analyses to characterize the spatiotemporal mechanisms of growth of new leaves and storage protein degradation in mature leaves of citrus trees exposed to low-N and high-N fertilization in the field. Results show that the growth of new leaves is driven by remobilization of stored reserves, rather than N uptake by the roots. In this context, proline and arginine in mature leaves acted as N sources supporting the growth of new leaves in spring. Time-series analyses with gel electrophoresis and proteome analysis indicated that the mature autumn shoot leaves are probably the sites of storage protein synthesis, while the aspartic endopeptidase protein is related to the degradation of storage proteins in mature citrus leaves. Furthermore, bioinformatic analysis based on protein-protein interactions indicated that glutamate synthetase and ATP-citrate synthetase are hub proteins in N remobilization from mature citrus leaves. These results provide strong physiological data for seasonal optimization of N fertilizer application in citrus orchards., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Interaction of different environmental constrains pose severe threats to plants that cannot be predicted from individual stress exposure. In this context, mercury (Hg), as a typical toxic and hazardous heavy metal, has recently attracted particular attention. Nitrogen (N 2 )-fixing legumes can be used for phytoremediation of Hg accumulation, whereas N availability could greatly affect its N 2 -fixation efficiency. However, information on the physiological responses to combined Hg exposure and excess N supply of woody legume species is still lacking. Here, we investigated the interactive effects of rhizobia inoculation, Hg exposure (+Hg), and high N (+N) supply, individually and in combination (+N*Hg), on photosynthesis and biochemical traits in Robinia pseudoacacia L. seedlings of two provenances, one from Northeast (DB) and one from Northwest (GS) China. Our results showed antagonistic effects of combined + N*Hg exposure compared to the individual treatments that were provenance-specific. Compared to individual Hg exposure, combined + N*Hg stress significantly increased foliar photosynthesis (+50.6%) of inoculated DB seedlings and resulted in more negative (-137.4%) δ 15 N abundance in the roots. Furthermore, combined + N*Hg stress showed 47.7% increase in amino acid N content, 39.4% increase in NR activity, and 14.8% decrease in MDA content in roots of inoculated GS seedlings. Inoculation with rhizobia significantly promoted Hg uptake in both provenances, reduced MDA contents of leaves and roots, enhanced photosynthesis and maintained the nutrient balance of Robinia. Among the two Robinia provenances investigated, DB seedlings formed more nodules, had higher biomass and Hg accumulation than GS seedlings. For example, total Hg concentrations in leaves and roots and total biomass of inoculated DB seedlings were 1.3,1.9 and 3.4 times higher than in inoculated GS seedlings under combined + N*Hg stress, respectively. Therefore, the DB provenance is considered to possess a higher potential for phytoremediation of Hg contamination compared to the GS provenance in environments subjected to N deposition., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Tropospheric ozone (O 3 ) is a significant phytotoxic air pollutant that has a negative impact on plant carbon gain. Although date palm (Phoenix dactylifera L.) is a globally important crop in arid or semi-arid regions, so far O 3 risk assessment for this species has not been reported. This study estimated leaf- and plant-level photosynthetic CO 2 uptake for understanding how elevated levels of O 3 affects date palm biomass growth. Ozone risks to date palm plants were assessed based on exposure- (AOT40) or flux-based indices (Phytotoxic Ozone Dose, POD y , where y is a threshold of uptake). For this purpose, plants were exposed to three levels of O 3 [ambient air, AA (45 ppb as daily average); 1.5 × AA; 2.0 × AA] for 92 days in an O 3 Free-Air Controlled Exposure facility. According to the model simulations, the negative effects of O 3 on plant-level net photosynthetic CO 2 uptake were attributed to reduced gross photosynthetic carbon gain and increased respiratory carbon loss. Season-long O 3 exposure and elevated temperatures promoted the negative O 3 effect because of a further increase of respiratory carbon loss, which was caused by increased leaf temperature due to stomatal closure. POD 1 nonlinearly affected the photosynthetic CO 2 uptake, which was closely related to the variation of dry mass increment during the experiment. Although the dose-response relationship suggested that a low O 3 dose (POD 1  < 5.2 mmol m -2 ) may even positively affect photosynthetic CO 2 uptake in date palms, stomatal O 3 uptake at the current ambient O 3 levels has potentially a negative impact on date palm growth. The results indicate 5.8 mmol m -2 POD 1 or 21.1 ppm h AOT40 as critical levels corresponding to a 4% reduction of net CO 2 uptake for date palm, suggesting that this species can be identified as a species moderately sensitive to O 3 ., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Urea is the most frequently used nitrogen (N) fertilizer worldwide. However, the mechanisms in plants to cope with excess urea are largely unknown, especially for woody legumes that can meet their N demand by their own N 2 -fixation capacity. Here, we studied the immediate consequences of different amounts of urea application and exposure duration on photosynthesis, N metabolism, and the activity of antioxidative enzymes of Robinia pseudoacacia seedlings. For this purpose, seedlings were grown for 3 months under normal N availability with rhizobia inoculation and, subsequently, 50 mg N kg -1 was applied to the soil twice with urea as additional N source. Our results show that excess urea application significantly promoted photosynthesis, which increased by 80.3% and 84.7% compared with CK after the 1st and 2nd urea applications, respectively. The increase in photosynthesis translated into an increase in root and nodule biomass of 88.7% and 82.0%, respectively, while leaf biomass decreased by 4.8% after the first application of urea. The N content in leaves was 92.6% higher than in roots, but excess urea application increased the N content of protein and free amino acids in roots by 25.0%, and 43.3%, respectively. Apparently, enhanced root growth and N storage in the roots constitute mechanisms to prevent the negative consequences of excess N in the shoot upon urea application. Nitrate reductase (NR) activity of leaves and roots increased by 74.4% and 26.3%, respectively. Glutathione reductase (GR) activity in leaves and roots was enhanced by 337% and 34.0%, respectively, but then decreased rapidly to the initial level before fertilization. This result shows that not only N metabolism, but also antioxidative capacity was transiently promoted by excess urea application. Apparently, excess urea application initially poses oxidative stress to the plants that is immediately counteracted by enhanced scavenging of reactive oxygen species via enhanced GR activity., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Anthropogenic activities such as mining, smelting, and overapplication of fertilizers contribute to introducing cadmium (Cd) into the biosphere. Cd accumulation in edible plants leads to phytotoxicity and reduces biomass formation and food production, posing a significant threat to global food security. Nitric oxide (NO) is a highly active gaseous signalling molecule involved in regulating plant responses to Cd stress. These responses include the protective role of NO in enhancing plant resistance to Cd exposure via activating the antioxidant defense system, maintaining intracellular redox homeostasis, and initiating the expression of genes relevant to stress defense. However, NO exacerbates Cd toxicity by promoting Cd uptake and accelerating programmed cell death in plants. These contradictory responses render the role of NO in regulating plant performance under Cd exposure highly controversial. To better understand the mechanisms responsible for the dual role of NO, we summarized the current knowledge on (1) the processes of Cd accumulation and detoxification in plants, (2) the pathways of NO synthesis and metabolism under Cd stress, and (3) the function of NO in regulating plant responses to Cd stress at the physiological and molecular levels. From this literature review, the processes responsible for the dual role of NO in plant responses to Cd exposure were deduced, and topics for future studies on the mechanisms of NO-mediated regulation of Cd detoxification in plants were identified., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

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., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Atmospheric nitrogen (N 2 )-fixing legume trees are frequently used for the restoration of depleted, degraded, and contaminated soils. However, biological N 2 fixation (BNF) can also be performed by so-called actinorhizal plants. Actinorhizal plants include a high diversity of woody species and therefore can be applied in a broad spectrum of environments. In contrast to N 2 -fixing legumes, the potential of actinorhizal plants for soil restoration remains largely unexplored. In this Opinion, we propose related basic research requirements for the characterization of environmental stress responses that determine the restoration potential of actinorhizal plants for depleted, degraded, and contaminated soils. We identify advantages and unexplored processes of actinorhizal plants and describe a mainly uncharted avenue of future research for this important group of plant species., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Rice consumption is an essential cause of mercury (Hg) exposure for humans in Asia. However, the mechanism of Hg transport and accumulation in rice plants (Oryza sativa L.) remains unclear. Here, rice genotypes with contrasting Hg uptake and translocation abilities, i.e. H655 (high Hg-accumulator) and H767 (low Hg-accumulator), were selected from 261 genotypes. Through comparative physiological and transcriptome analyses, we investigated the processes responsible for the relationship between Hg accumulation, transport and tolerance. The results showed significant stimulation of antioxidative metabolism, particularly glutathione (GSH) accumulation, and up-regulated expression of regulatory genes of glutathione metabolism for H655, but not for H767. In addition, up-regulated expression of GSH S-transferase (GST) and OsPCS1 in H655 that catalyzes the binding of Hg and GSH, enhances the Hg detoxification capacity, while high-level expression of YSL2 in H655 enhances the transport ability for Hg. Conclusively, Hg accumulation in rice is a consequence of enhanced expression of genes related to Hg binding with GSH and Hg transport. With these results, the present study contributes to the selection of rice genotypes with limited Hg accumulation and to the mitigation of Hg migration in food chains thereby enhancing nutritional safety of Hg-polluted rice fields., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Infection with the necrotrophic fungus Diplodia sapinea (Fr.) Fuckel is among the economically and ecologically most devastating diseases of conifers in the northern hemisphere and is accelerated by global climate change. This study aims to characterize the changes mediated by D. sapinea infection on its pine host (Pinus sylvestris L.) that lead to the death of its needles. For this purpose, we performed an indoor infection experiment and inoculated shoot tips of pine seedlings with virulent D. sapinea. The consequences for foliar traits, including the phytohormone profile, were characterized at both the metabolite and transcriptome level. Our results showed that D. sapinea infection strongly affected foliar levels of most phytohormones and impaired a multitude of other metabolic and structural foliar traits, such as reactive oxygen species scavenging. Transcriptome analysis revealed that these changes are partially mediated via modified gene expression by fungal exposure. Diplodia sapinea appears to overcome the defense reactions of its pine host by reprogramming gene expression and post-transcriptional controls that determine essential foliar metabolic traits such as the phytohormone profile, cell wall composition and antioxidative system., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Drought and salt exposure are among the most prevalent and severe abiotic stressors causing serious agricultural yield losses, alone and in combination. Little is known about differences and similarities in the effects of these two stress factors on plant metabolic regulation, particularly on nitrogen metabolism. Here, we studied the effects of water deprivation and salt exposure on water relations and nitrogen metabolites in leaves and roots of date palm seedlings. Both, water deprivation and salt exposure had no significant effects on plant water content or stable carbon (C) and nitrogen (N) isotope signatures. Significant effects of water deprivation on total C and N concentrations were only observed in roots, i.e., decreased total C and increased total N concentrations. Whereas salt exposure initially decreased total C and increased total N concentrations significantly in roots, foliar total C concentration was increased upon prolonged exposure. Initially C/N ratios declined in roots of plants from both treatments and upon prolonged salt exposure also in the leaves. Neither treatment affected soluble protein and structural N concentrations in leaves or roots, but resulted in the accumulation of most amino acids, except for glutamate and tryptophan, which remained stable, and serine, which decreased, in roots. Accumulation of the most abundant amino acids, lysine and proline, was observed in roots under both treatments, but in leaves only upon salt exposure. This finding indicates a similar role of these amino acids as compatible solutes in the roots in response to salt und drought, but not in the leaves. Upon prolonged treatment, amino acid concentrations returned to levels found in unstressed plants in leaves of water deprived, but not salt exposed, plants. The present results show both water deprivation and salt exposure strongly impact N metabolism of date palm seedlings, but in a different manner in leaves and roots., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

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., (© 2023. The Author(s).)

Chronic ozone (O 3 ) exposure in the atmosphere preferentially disturbs metabolic processes in the roots rather than the shoot as a consequence of reduced photosynthesis and carbohydrate allocation from the leaves to the roots. The aim of the present study was to elucidate if mineral nutrition is also impaired by chronic O 3 exposure. For this purpose, date palm (Phoenix dactylifera) plants were fumigated with ambient, 1.5 × ambient and 2 × ambient O 3 in a free air controlled exposure (FACE) system for one growing season and concentrations of major nutrients were analyzed in leaves and roots. In addition, concentrations of C and N and their partitioning between different metabolic C and N pools were determined in both organs. The results showed that calcium (Ca), magnesium (Mg), iron (Fe), zinc (Zn), sodium (Na) and potassium (K) acquisition by roots was diminished by O 3 exposure of the shoot. For Ca, Mg, Fe and Zn reduced uptake by the roots was combined with reduced allocation to the shoot, resulting in a decline of foliar concentrations; for Na and K, allocation to the shoot was maintained at the expense of the roots. Thus, elevated O 3 impaired both mineral uptake by the roots and partitioning of minerals between roots and shoots, but in an element specific way. Thereby, elevated O 3 affected roots and shoots differently already after one growing season. However, considerable changes in total C and N concentrations and their partitioning between different metabolic pools upon chronic O 3 exposure were not observed in either leaves or roots, except for reduced foliar lignin concentrations at 2 × ambient O 3 . Significant differences in these parameters were shown between leaves and roots independent of O 3 application. The physiological consequences of the effects of chronic O 3 exposure on mineral acquisition and partitioning between leaves and roots are discussed., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Nitrogen (N) remobilization is an important physiological process that supports the growth and development of trees. However, in evergreen broad-leaved tree species, such as citrus, the mechanisms of N remobilization are not completely understood. Therefore, we quantified the potential of N remobilization from senescing leaves of spring shoots to mature leaves of autumn shoots of citrus trees under different soil N availabilities and further explored the underlying N metabolism characteristics by physiological, proteome and gene expression analyses. Citrus exposed to low N had an approximately 38% N remobilization efficiency (NRE), whereas citrus exposed to high N had an NRE efficiency of only 4.8%. Integrated physiological, proteomic and gene expression analyses showed that photosynthesis, N and carbohydrate metabolism interact with N remobilization. The improvement of N metabolism and photosynthesis, the accumulation of proline and arginine, and delayed degradation of storage protein in senescing leaves are the result of sufficient N supply and low N remobilization. Proteome further showed that energy generation proteins and glutamate synthase were hub proteins affecting N remobilization. In addition, N requirement of mature leaves is likely met by soil supply at high N nutrition, thereby resulting in low N remobilization. These results provide insight into N remobilization mechanisms of citrus that are of significance for N fertilizer management in orchards., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Selenium-binding proteins (SBPs) represent a ubiquitous protein family implicated in various environmental stress responses, although the exact molecular and physiological role of the SBP family remains elusive. In this work, we report the identification and characterization of CrSBD1, an SBP homolog from the model microalgae Chlamydomonas reinhardtii. Growth analysis of the C. reinhardtii sbd1 mutant strain revealed that the absence of a functional CrSBD1 resulted in increased growth under mild oxidative stress conditions, although cell viability rapidly declined at higher hydrogen peroxide (H2O2) concentrations. Furthermore, a combined global transcriptomic and metabolomic analysis indicated that the sbd1 mutant exhibited a dramatic quenching of the molecular and biochemical responses upon H2O2-induced oxidative stress when compared to the wild-type. Our results indicate that CrSBD1 represents a cell regulator, which is involved in the modulation of C. reinhardtii early responses to oxidative stress. We assert that CrSBD1 acts as a member of an extensive and conserved protein-protein interaction network including Fructose-bisphosphate aldolase 3, Cysteine endopeptidase 2, and Glutaredoxin 6 proteins, as indicated by yeast two-hybrid assays., (© American Society of Plant Biologists 2022. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Karst lime soil, commonly found in rocky desert ecosystems of Southwest China, exhibits high pH, poor water retention, and intense erosion. To prevent further soil erosion and soil losses from these ecosystems, stabilization measures based on improved green infrastructure are needed. The present study aimed at elucidating the performance of sweet tea (Lithocarpus polystachyus) seedlings grown on this soil type upon biochar application. Biochar was classified into different particle sizes, viz. 0.25-0.5 mm (medium), 0.5-1 mm (coarse), 1-2 mm (gravel), and their mixture, and added at the concentrations of 1, 2, or 5% soil mass. The pH, moisture, and porosity of soil increased upon biochar application compared to control; however, soil bulk density significantly decreased. The activity of soil phosphatase was increased by biochar particle size. Biochar particle size and concentration significantly enhanced the soil organic carbon content, but they differently affected total and plant-available nutrients in the soil. Light-saturated photosynthesis was positively affected, while stomatal conductance, leaf transpiration, and the intercellular CO 2 concentrations of sweet tea leaves were negatively affected by biochar particle size and/or concentration compared to control. Leaf chlorophyll and soluble protein contents were increased by biochar application. From these results, we conclude that biochar can improve soil properties and the performance of sweet tea seedlings grown on Karst lime soil. We suggest its application at a concentration of 2% soil mass for keeping a high physiological performance of sweet tea seedlings in this environment. The selection of the ideal particle size is context-specific and depends on the target outcome., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The microbe-clay mineral system is widely known to reduce the fluidity of heavy metals through biomineralization, thus mitigating soil pollution stemming from heavy metals. Here, we investigated the effect of mineral distinction on the solidification of cadmium (Cd) using sulfate-reducing bacteria (SRB) to construct symbiotic systems with purplish soil, clay-sized fraction of purple soil (Clay -csp ), clay particles of amorphous iron (Fe) oxide (Clay -ox ), clay particles removing crystalline Fe oxide (Clay -CBD ), and residues of Clay -CBD treated by hydrochloric acid (Clay -HCl ). The difference in Cd morphology among purplish soil, Clay -csp , and Clay -ox indicated that the fixation of Cd in soil was largely determined by Fe oxides. The content of Cd in Clay -csp decreased by 66.7% after the removal of amorphous Fe, confirming that clay easily adsorbed infinitive Fe oxides in purple soil. In the system of SRB and Clay -ox , carbonate-bound Cd (F2) decreased by 14.85% and residual Cd (F5) increased by 14% from the retardation to late decline phase, eventually forming iron-sulfur (Fe-S) compounds. Based on the correlation analyses of Cd and Fe in amorphous-bound state and Fe-manganese (Mn) oxidation state in simulation experiments, it is demonstrated that Fe-Mn oxides control the behavior of Cd in soil clay, and SRB-mediated Fe-bearing minerals promote the transformation of Cd from activated to stable state., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The present study explores the interaction of water supply and rhizobia inoculation on CO 2 and H 2 O gas exchange characteristics, physiological and biochemical traits in seedlings of Robinia pseudoacacia L. originating from two provenances with contrasting climate and soil backgrounds: the Gansu Province (GS) in northwest China and the Dongbei region (DB) of northeast China. Rhizobia strains were isolated from the 50-years old Robinia forest sites grown in the coastal region of east China. Robinia seedlings with and without rhizobia inoculation were exposed to normal water supply, moderate drought, and rewatering treatments, respectively. After 2 weeks of drought treatment, photosynthetic and physiological traits (net photosynthetic rate, stomatal conductance, stable isotope signature of carbon, malondialdehyde and hydrogen peroxide content) of Robinia leaves were significantly altered, but after rewatering, a general recovery was observed. Rhizobia inoculation significantly increased the drought resistance of both Robinia provenances by promoting photosynthesis, increasing the foliar N content and reducing the accumulation of malondialdehyde and hydrogen peroxide. Among the two provenances, DB plants developed more nodules than GS plants, but GS plants were more drought-tolerant than DB plants, both inoculated or noninoculated, indicated by the foliar gas exchange parameters and biochemical traits studied. Our results also show that inoculation of rhizobia could significantly improve the drought resistance of Robinia in both provenances. The present study contributes to the scientific background for the selection of drought-resistant varieties of Robinia to ensure the success of future afforestation projects in degraded terrestrial ecosystems under global climate change., (© 2022 Scandinavian Plant Physiology Society.)

The spring shoot leaves are important sites of nitrogen (N) metabolism in citrus trees. Understanding the physiological and metabolic response of the spring shoot leaves under varying N fertilization is fundamental to the fertilization management in citrus orchards. Thus, the processes affecting N composition, the activities of N metabolism related enzymes, and the expression of relevant genes were explored in spring shoot leaves under four N levels (0, 207, 275, 413 g N tree -1 y -1 , as N0, N207, N275, N413). The results showed that, compared with N0, N275 significantly increased total N by 24.81%, which was mainly attributed to enhancement of structural N by 30.92%, free amino acid N by 40.91% and nitrate N by 41.33%. The relative expression of nitrate reductase (NR) and glutamate dehydrogenase (GDH) under N275 increased by 19.32% and 73.48%, respectively, compared with that under N0 treatment. Compared with N0 treatment, the NR transcription level under N275 treatment increased by 381%. The relative transcription levels of NADP-GDH and GDH1 also increased with increasing N fertilization. However, compared with that under N275, the relative transcription of GDH2 under N413 treatment was inhibited. Therefore, the transcript abundance of NR, NADP-GDH,GDH1 and GDH2 affected the activities of NR and GDH and thereby contributed to the regulation of N composition in the leaves. In addition, the activities of glutamine synthetase and nitrite reductase were largely unaffected or even declined in the N207, N275 and N413 treatments compared with the N0. This study elucidated the mechanism of primary N metabolism and partitioning in citrus leaves and provided a theoretical basis for N management in citrus orchards., (Copyright © 2021 Elsevier GmbH. All rights reserved.)

Climate change is increasing the frequency and intensity of warming and drought periods around the globe, currently representing a threat to many plant species. Understanding the resistance and resilience of plants to climate change is, therefore, urgently needed. As date palm (Phoenix dactylifera) evolved adaptation mechanisms to a xeric environment and can tolerate large diurnal and seasonal temperature fluctuations, we studied the protein expression changes in leaves, volatile organic compound emissions, and photosynthesis in response to variable growth temperatures and soil water deprivation. Plants were grown under controlled environmental conditions of simulated Saudi Arabian summer and winter climates challenged with drought stress. We show that date palm is able to counteract the harsh conditions of the Arabian Peninsula by adjusting the abundances of proteins related to the photosynthetic machinery, abiotic stress and secondary metabolism. Under summer climate and water deprivation, these adjustments included efficient protein expression response mediated by heat shock proteins and the antioxidant system to counteract reactive oxygen species formation. Proteins related to secondary metabolism were downregulated, except for the P. dactylifera isoprene synthase (PdIspS), which was strongly upregulated in response to summer climate and drought. This study reports, for the first time, the identification and functional characterization of the gene encoding for PdIspS, allowing future analysis of isoprene functions in date palm under extreme environments. Overall, the current study shows that reprogramming of the leaf protein profiles confers the date palm heat- and drought tolerance. We conclude that the protein plasticity of date palm is an important mechanism of molecular adaptation to environmental fluctuations., (© 2021. The Author(s).)