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Soil salinization has been considered as a major environmental threat to plant growth. Different types of salt in saline soil have different effects on germination and seedling growth. Effect of NaCl on germination and seedling establishment in Suaeda liaotungensis have been reported, but its response to alkali stress remains unclear. Our results showed that brown seeds had higher germination rate, however, black seeds had higher germination recovery percentage under alkali stress. Na 2 CO 3 had stronger inhibitory effect on germination and seedling growth than NaHCO 3 . As the concentration of alkali stress increased, the ROS level of brown seeds gradually ascended, while that of black seeds decreased first and then ascended. MDA content of dimorphic seeds significantly increased under alkali stress. The trend of SOD, POD and CAT activity between dimorphic seeds was similar under the same type of alkali stress. Alkali stress enhanced proline content of dimorphic seeds, and dimorphic seeds in NaHCO 3 solution had higher proline content than Na 2 CO 3 solution. Moreover, radicle and shoot tolerance indexes of seedlings in NaHCO 3 solution were significantly higher than that of Na 2 CO 3 solution. Under strong alkali stress, seedlings in NaHCO 3 solution had significantly lower ROS level and MDA content as well as higher antioxidant enzyme activity than Na 2 CO 3 solution. This study comprehensively compared the morphological and physiological characteristics in germination and seedlings to better reveal the saline-alkali tolerance mechanisms in S. liaotungensis., (© 2024. The Author(s) under exclusive licence to The Botanical Society of Japan.)

Soil salinity is a significant abiotic stress and poses risks to environmental sustainability. Thus, the improvement of the time for recovering the salt-affect soil is crucial for the phytoextraction process using halophytes plants, especially regarding on nutritional management. We evaluated the responses of Atriplex nummularia Lindl. to nitrogen (N) and phosphorus (P) under different salinity levels. The treatments comprised doses of N (N1 = 80 kg ha -1 ) and P (P1 = 60 kg ha -1 ): (1) without N and P (N0P0) (control); (2) with N and without P (N1P0); (3) without N and with P (N0P1); and (4) with N and P (N1P1) and five levels of electrical conductivity from irrigation water: 0.08, 1.7, 4.8, 8.6, and 12.5 dS m -1 . The. We evaluated dry biomass of leaves, stems, and roots 93 days after transplantation. We also assessed the leaf and osmotic water potential, the osmotic adjustment, and the nutrient contents (N, P, Na, and K). N application increased 22.3, 17.8, and 32.8% the leaf biomass, stem biomass, and osmotic adjustment, respectively; and consequently, boosts Na extraction in 27.8%. Thus, the time of the phytoextraction process can be improved with N fertilizer at a rate of 80 kg ha -1 .

Climate and edaphic properties drive the biogeographic distribution of dominant soil microbial phylotypes in terrestrial ecosystems. However, the impact of plant species and their root nutritional traits on microbial distribution in coastal wetlands remains unclear. Here, we investigated the nutritional traits of 100 halophyte root samples and the bacterial communities in the corresponding soil samples from coastal wetlands across eastern China. This study spans 22° of latitude, covering over 2500 km from north to south. We found that 1% of soil bacterial phylotypes accounted for nearly 30% of the soil bacterial community abundance, suggesting that a few bacterial phylotypes dominated the coastal wetlands. These dominated phylotypes could be grouped into three ecological clusters as per their preference over climatic (temperature and precipitation), edaphic (soil carbon and nitrogen), and plant factors (halophyte vegetation, root carbon, and nitrogen). We further provide novel evidence that plant root nutritional traits, especially root C and N, can strongly influence the distribution of these ecological clusters. Taken together, our study provides solid evidence of revealing the dominance of specific bacterial phylotypes and the complex interactions with their environment, highlighting the importance of plant root nutritional traits on biogeographic distribution of soil microbiome in coastal wetland ecosystems., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

The escalating salinity levels in cultivable soil pose a significant threat to agricultural productivity and, consequently, human sustenance. This problem is being exacerbated by natural processes and human activities, coinciding with a period of rapid population growth. Developing halophytic crops is needed to ensure food security is not impaired and land resources can be used sustainably. Evolution has created many close halophyte relatives of our major glycophytic crops, such as Puccinellia tenuiflora (relative of barley and wheat), Oryza coarctata (relative of rice) and Glycine soja (relative of soybean). There are also some halophytes have been subjected to semi-domestication and are considered as minor crops, such as Chenopodium quinoa. In this paper, we examine the prevailing comprehension of robust salinity resilience in halophytes. We summarize the existing strategies and technologies that equip researchers with the means to enhance the salt tolerance capabilities of primary crops and investigate the genetic makeup of halophytes., Competing Interests: Declaration of Competing Interest No conflict of interest exits in the submission of this manuscript, and manuscript is approved by all authors for publication. I would like to declare on behalf of my co-authors that the work described was original research that has not been published previously, and not under consideration for publication elsewhere, in whole or in part. All the authors listed have approved the manuscript that is enclosed., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Soil salinization leads to reduced crop yields and waste of land resources, thereby impacting global food security. To meet the increasing demand for food and simultaneously alleviate pressure on soil resources, the development of sustainable agriculture is imperative. In contrast to physical and chemical methods, bioremediation represents an efficient and environmentally friendly approach. Fungal symbionts have been found to be associated with most plants in natural ecosystems, colonizing and residing within the internal tissues of host plants. Moreover, the potential of fungal symbionts in improving saline-alkaline soil has been widely recognized and confirmed. Numerous reports have documented the effectiveness of arbuscular mycorrhizal fungi in alleviating salt stress in plants. Meanwhile, research on other endophytic fungi for mitigating plant salt stress has emerged in recent years, which contributes to refining mechanisms for enhancing plant salt tolerance. In this review, we summarized various mechanisms by which endophytic fungi enhance plant salt tolerance. We also provided an overview of the challenges and development directions in the field of fungal symbiosis, with the aim of offering a viable strategy for the bioremediation of saline-alkali soils., 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.)

Background: Suaeda australis is one of typical halophyte owing to high levels of salt tolerance. In addition, the bZIP gene family assumes pivotal functions in response to salt stress. However, there are little reports available regarding the bZIP gene family in S. australis., Results: In this study, we successfully screened 44 bZIP genes within S. australis genome. Subsequently, we conducted an extensive analysis, encompassing investigations into chromosome location, gene structure, phylogenetic relationship, promoter region, conserved motif, and gene expression profile. The 44 bZIP genes were categorized into 12 distinct groups, exhibiting an uneven distribution among the 9 chromosomes of S. australis chromosomes, but one member (Sau23745) was mapped on unanchored scaffolds. Examination of cis-regulatory elements revealed that bZIP promoters were closely related to anaerobic induction, transcription start, and light responsiveness. Comparative transcriptome analysis between ST1 and ST2 samples identified 2,434 DEGs, which were significantly enriched in some primary biological pathways related to salt response-regulating signaling based on GO and KEGG enrichment analysis. Expression patterns analyses clearly discovered the role of several differently expressed SabZIPs, including Sau08107, Sau08911, Sau11415, Sau16575, and Sau19276, which showed higher expression levels in higher salt concentration than low concentration and a response to salt stress. These expression patterns were corroborated through RT-qPCR analysis. The six differentially expressed SabZIP genes, all localized in the nucleus, exhibited positive regulation involved in the salt stress response. SabZIP14, SabZIP26, and SabZIP36 proteins could bind to the promoter region of downstream salt stress-related genes and activate their expressions., Conclusions: Our findings offer valuable insights into the evolutionary trajectory of the bZIP gene family in S. australis and shed light on their roles in responding to salt stress. In addition to fundamental genomic information, these results would serve as a foundational framework for future investigations into the regulation of salt stress responses in S. australis., (© 2024. The Author(s).)

Mangroves perform a crucial ecological role along the tropical and subtropical coastal intertidal zone where salinity fluctuation occurs frequently. However, the differential responses of mangrove plant at the combined transcriptome and metabolome level to variable salinity are not well documented. In this study, we used Avicennia marina (Forssk.) Vierh., a pioneer species of mangrove wetlands and one of the most salt-tolerant mangroves, to investigate the differential salt tolerance mechanisms under low and high salinity using inductively coupled plasma-mass spectrometry, transcriptomic and metabolomic analysis. The results showed that HAK8 was up-regulated and transported K+ into the roots under low salinity. However, under high salinity, AKT1 and NHX2 were strongly induced, which indicated the transport of K+ and Na+ compartmentalization to maintain ion homeostasis. In addition, A. marina tolerates low salinity by up-regulating ABA signaling pathway and accumulating more mannitol, unsaturated fatty acids, amino acids' and L-ascorbic acid in the roots. Under high salinity, A. marina undergoes a more drastic metabolic network rearrangement in the roots, such as more L-ascorbic acid and oxiglutatione were up-regulated, while carbohydrates, lipids and amino acids were down-regulated in the roots, and, finally, glycolysis and TCA cycle were promoted to provide more energy to improve salt tolerance. Our findings suggest that the major salt tolerance traits in A. marina can be attributed to complex regulatory and signaling mechanisms, and show significant differences between low and high salinity., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Premise: The origin of diversity is a fundamental biological question. Gene duplications are one mechanism that provides raw material for the emergence of novel traits, but evolutionary outcomes depend on which genes are retained and how they become functionalized. Yet, following different duplication types (polyploidy and tandem duplication), the events driving gene retention and functionalization remain poorly understood. Here we used Cakile maritima, a species that is tolerant to salt and heavy metals and shares an ancient whole-genome triplication with closely related salt-sensitive mustard crops (Brassica), as a model to explore the evolution of abiotic stress tolerance following polyploidy., Methods: Using a combination of ionomics, free amino acid profiling, and comparative genomics, we characterize aspects of salt stress response in C. maritima and identify retained duplicate genes that have likely enabled adaptation to salt and mild levels of cadmium., Results: Cakile maritima is tolerant to both cadmium and salt treatments through uptake of cadmium in the roots. Proline constitutes greater than 30% of the free amino acid pool in C. maritima and likely contributes to abiotic stress tolerance. We find duplicated gene families are enriched in metabolic and transport processes and identify key transport genes that may be involved in C. maritima abiotic stress tolerance., Conclusions: These findings identify pathways and genes that could be used to enhance plant resilience and provide a putative understanding of the roles of duplication types and retention on the evolution of abiotic stress response., (© 2024 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Background: Alfalfa (Medicago sativa L.) experiences many negative effects under salinity stress, which may be mediated by recurrent selection. Salt-tolerant alfalfa may display unique adaptations in association with rhizobium under salt stress., Results: To elucidate inoculation effects on salt-tolerant alfalfa under salt stress, this study leveraged a salt-tolerant alfalfa population selected through two cycles of recurrent selection under high salt stress. After experiencing 120-day salt stress, mRNA was extracted from 8 random genotypes either grown in 0 or 8 dS/m salt stress with or without inoculation by Ensifer meliloti. Results showed 320 and 176 differentially expressed genes (DEGs) modulated in response to salinity stress or inoculation x salinity stress, respectively. Notable results in plants under 8 dS/m stress included upregulation of a key gene involved in the Target of Rapamycin (TOR) signaling pathway with a concomitant decrease in expression of the SNrK pathway. Inoculation of salt-stressed plants stimulated increased transcription of a sulfate-uptake gene as well as upregulation of the Lysine-27-trimethyltransferase (EZH2), Histone 3 (H3), and argonaute (AGO, a component of miRISC silencing complexes) genes related to epigenetic and post-transcriptional gene control., Conclusions: Salt-tolerant alfalfa may benefit from improved activity of TOR and decreased activity of SNrK1 in salt stress, while inoculation by rhizobiumstimulates production of sulfate uptake- and other unique genes., (© 2024. The Author(s).)

The use of plant growth-promoting rhizobacteria (PGPR) in agriculture is one of the most promising approaches to improve plants' growth under salt stress and to support sustainable agriculture under climate change. In this context, our goal was to grow and enhance quinoa growth using native rhizobacteria that can withstand salt stress. To achieve this objective, we isolated rhizobacteria from three saline localities in a semi-arid region in Tunisia, which are characterized by different halophyte species and tested their plant growth-promoting (PGP) activities. Then, we inoculated quinoa seedlings cultivated on 300 mM NaCl with the three most efficient rhizobacteria. A positive effect of the three-salt tolerant rhizobacteria on the growth of quinoa under salinity was observed. In fact, the results of principal component analysis indicated that the inoculation of quinoa by salt-tolerant PGPR under high salinity had a prominent beneficial effect on various growth and physiological parameters of stressed plant, such as the biomass production, the roots length, the secondary roots number, proline content and photosynthesis activities. Three rhizobacteria were utilized in this investigation, and the molecular identification revealed that strain 1 is related to the Bacillus inaquosorum species, strain 2 to Bacillus thuringiensis species and strain 3 to Bacillus proteolyticus species. We can conclude that the saline soil, especially the halophytic rhizosphere, is a potential source of salt-tolerant plant growth-promoting rhizobacteria (ST-PGPR), which stimulate the growth of quinoa and improve its tolerance to salinity., (© 2024 Scandinavian Plant Physiology Society.)

Background and Aims: Seed heteromorphism is a plant strategy that an individual plant produces two or more distinct types of diaspores, which have diverse morphology, dispersal ability, ecological functions and different effects on plant life history traits. The aim of this study was to test the effects of seasonal soil salinity and burial depth on the dynamics of dormancy/germination and persistence/depletion of buried trimorphic diaspores of a desert annual halophyte Atriplex centralasiatica., Methods: We investigated the effects of salinity and seasonal fluctuations of temperature on germination, recovery of germination and mortality of types A, B, C diaspores of A. centralasiatica in the laboratory and buried diaspores in situ at four soil salinities and three depths. Diaspores were collected monthly from the seedbank from December 2016 to November 2018, and the number of viable diaspores remaining (not depleted) and their germinability were determined., Results: Non-dormant type A diaspores were depleted in the low salinity "window" in the first year. Dormant diaspore types B and C germinated to high percentages at 0.3 and 0.1 mol L -1 soil salinity, respectively. High salinity and shallow burial delayed depletion of diaspore types B and C. High salinity delayed depletion time of the three diaspore types and delayed dormancy release of types B and C diaspores from autumn to spring. Soil salinity modified the response of diaspores in the seedbank by delaying seed dormancy release in autum and winter and by providing a low-salt concentration window for germination of non-dormant diaspores in spring and early summer., Conclusions: Buried trimorphic diaspores of annual desert halophyte A. centralasiatica exhibited diverse dormancy/germination behavior in respond to seasonal soil salinity fluctuation. Prolonging persistence of the seedbank and delaying depletion of diaspores under salt stress in situ primarily is due to inhibition of dormancy-break. The differences in dormancy/germination and seed persistence in the soil seedbank may be a bet-hadging strategy adapted to stressful temporal and spatial heterogeneity, and allows A. centralasiatica to persist in the unpredictable cold desert enevironment., (© 2024. The Author(s).)

Halophyte Halogeton glomeratus mostly grows in saline desert areas in arid and semi-arid regions and is able to adapt to adverse conditions such as salinity and drought. Earlier transcriptomic studies revealed activation of the HgS2 gene in the leaf of H. glomeratus seedlings when exposed to saline conditions. To identify the properties of HgS2 in H. glomeratus, we used yeast transformation and overexpression in Arabidopsis. Yeast cells genetically transformed with HgS2 exhibited K + uptake and Na + efflux compared with control (empty vector). Stable overexpression of HgS2 in Arabidopsis improved its resistance to salt stress and led to a notable rise in seed germination in salinity conditions compared to the wild type (WT). Transgenic Arabidopsis regulated ion homeostasis in plant cells by increasing Na + absorption and decreasing K + efflux in leaves, while reducing Na + absorption and K + efflux in roots. In addition, overexpression of HgS2 altered transcription levels of stress response genes and regulated different metabolic pathways in roots and leaves of Arabidopsis. These results offer new insights into the role of HgS2 in plants' salt tolerance., (© 2024 Scandinavian Plant Physiology Society.)

Rice production is seriously affected by saline-alkaline stress worldwide. To elucidate the saline-alkaline tolerance mechanisms in a novel tolerant rice variety, Shwe Nang Gyi (SNG), we investigated ion accumulation in SNG and Koshihikari (KSH), which is a saline-alkaline sensitive rice variety, and the candidates for saline-alkaline inducible genes in SNG using RNA-seq. SNG had superior ion accumulation capacity, such as K and Zn, compared to KSH. In contrast, SNG accumulated the same level of Na content in its leaf blades as KSH despite the higher dry weight of the SNG leaf blades. We further found that the expression of numerous genes, including several K + transporter/high-affinity K + transporter/K + uptake protein/K + transporter (HAK/KUP/KT) family members, were upregulated in SNG, and that OsHAK17 and OsHAK21 expression levels in the roots were significantly higher in SNG than in KSH. Moreover, yeast complementation analysis revealed that OsHAK17 was involved in K + uptake under high-Na conditions. These results suggested that SNG has an effective K + acquisition system supported by OsHAK17 functioning in saline-alkaline environments., (© 2024. The Author(s).)

A huge amount of phosphogypsum (PG) wastes generated from the processing phosphate ore in Tunisia Industrial Group Area-Gabes is getting discarded into the sea. Within this framework, the basic objective of this research is to elaborate and discuss a natural-based solution focused on phytoremediation of contaminated (PG) soils and marine sediments with the halophilic plant Salicornia europaea. A significant drop of the organic matter (53.09%), moisture (26.47%), and sediment porosity with (5.88%) was detected in the rhizosphere Salicornia europaea area (RS). Removal of hazardous elements concentrations, such as Pb, Fe, Cu, Cd, and Zn, between contaminated sediment (CS) and RS displayed a significant difference, ranging from 5.33 to 50.02% of hazardous elements removal concentration, which was observed in the rhizosphere zone. The microbiota of both areas (RS and CS) were analyzed by massive sequencing. In both samples, all the sequences belong to only four phyla: Firmicutes and, to a much lower extent, Proteobacteria, Bacteroidetes, and Actinobacteria. The CS sediment seems to be heavily polluted by human activities. Most of the found genera are inhabitants of the intestine of warm-blooded animals (Escherichia, Bacteroides, Prevotella, Faecalibacterium, Ruminococcus, Enterococcus); hence, activities in this area pose a health risk. On the other hand, it may be surprising that 76.4% of the total high-quality sequences retrieved from the RS sample were affiliated to the family Bacillaceae. The salinity of the studied soil exerts a stress on the microbial populations that inhabit it, directing the selection of halotolerant species., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Salicylic acid (SA) is known to enhance salt tolerance in plants. However, the mechanism of SA-mediated response to high salinity in halophyte remains unclear. Using electrophysiological and molecular biological methods, we investigated the role of SA in response to high salinity in mangrove species, Kandelia obovata, a typical halophyte. Exposure of K. obovata roots to high salinity resulted in a rapid increase in endogenous SA produced by phenylalanine ammonia lyase pathway. The application of exogenous SA improved the salt tolerance of K. obovata, which depended on the NADPH oxidase-mediated H 2 O 2 . Exogenous SA and H 2 O 2 increased Na + efflux and reduced K + loss by regulating the transcription levels of Na + and K + transport-related genes, thus reducing the Na + /K + ratio in the salt-treated K. obovata roots. In addition, exogenous SA-enhanced antioxidant enzyme activity and its transcripts, and the expressions of four genes related to AsA-GSH cycle as well, then alleviated oxidative damages in the salt-treated K. obovata roots. However, the above effects of SA could be reversed by diphenyleneiodonium chloride (the NADPH oxidase inhibitor) and paclobutrazol (a SA biosynthesis inhibitor). Collectively, our results demonstrated that SA-induced salt tolerance of K. obovata depends on NADPH oxidase-generated H 2 O 2 that affects Na + /K + and redox homeostasis in response to high salinity., (© 2024 Society for Experimental Biology and John Wiley & Sons Ltd.)

Salt excretory halophytes are the major sources of phytoremediation of salt-affected soils. Cressa cretica is a widely distributed halophyte in hypersaline lands in the Cholistan Desert. Therefore, identification of key physio-anatomical traits related to phytoremediation in differently adapted C. cretica populations was focused on. Four naturally adapted ecotypes of non-succulent halophyte Cressa cretica L. form hyper-arid and saline desert Cholistan. The selected ecotypes were: Derawar Fort (DWF, ECe 20.8 dS m -1 ) from least saline site, Traway Wala Toba (TWT, ECe 33.2 dS m -1 ) and Bailah Wala Dahar (BWD, ECe 45.4 dS m -1 ) ecotypes were from moderately saline sites, and Pati Sir (PAS, ECe 52.4 dS m -1 ) was collected from the highly saline site. The natural population of this species was collected and carefully brought to the laboratory for different structural and functional traits. As a result of high salinity, Na + , Cl - , K + , and Ca 2+ content significantly increased at root and shoot level. At root level, some distinctive modifications such as increased sclerification in vascular bundles, enlarged vascular bundles, metaxylem vessels, phloem region, and storage parenchyma (cortex) are pivotal for water storage under extreme arid and osmotic condition. At the stem level, enhanced sclerification in outer cortex and vascular bundles, stem cellular area, cortical proportion, metaxylem and phloem area, and at the leaf level, very prominent structural adaptations were thicker and smaller leaves with increased density of salt glands and trichomes at surface, few and large stomata, reduced cortical and mesophyll parenchyma, and narrow xylem vessels and phloem area represent their non-succulent nature. The ecotype collected from hypersaline environments was better adapted regarding growth traits, ion uptake and excretion, succulence, and phytoremediation traits. More importantly, structural and functional traits such as root length and biomass, accumulation of toxic ions along with K + in root and shoot, accumulation of Ca 2+ in shoot and Mg 2+ in root, excretion of toxic ions were the highest in this ecotype. In conclusion, all these alterations strongly favor water conservation, which certainly contributes to ecotypes survival under salt-induced physiological drought.

This paper presents new data on the salt tolerance and avoidance mechanisms among various groups of halophytes in India. The halophytic flora in general has positive effect of high saline environments on growth and physiology. The coastal area of the Kachchh district in Gujarat includes about 350 km of shoreline along the Gulf of Kachchh. This study presents data on the elemental accumulation mechanisms in soil and halophytic flora (succulent and non-succulents). The halophytes were divided into two groups namely succulent with thick and fleshy leaves and stems as well as non-succulents with thin leaves and stem. The succulent halophytes included species such as Salicornia brachiata, Suaeda fruticosa and Suaeda nudiflora. The non-succulent halophytes include Aeluropus lagopoides and Urochondra setulosa. Plant parts namely leaves (Phylloclade for Salicornia), stems and roots were analyzed during the monsoon season. The results of soil and plant mineral ion contents differed widely across the intertidal zones in the same habitat. Likewise, the intra species have varied in all nutrient levels and salt concentration. The accumulation of elemental concentration was high during the monsoon season in the succulent Salicornia brachiata, especially in leaves that showed Na + reaching high up to 7.6 meq g -1 , whereas Cl - was noted to be 4.34 meq g -1 . In the non-succulent halophytes, the accumulation of mineral ion concentration was lower when compared to succulent plants., (© 2023. Springer Nature Limited.)

Saline agriculture may contribute to food production in the face of the declining availability of fresh water and an expanding area of salinized soils worldwide. However, there is currently little known about the biomass and nutrient/antinutrient accumulation response of many edible halophytes to increasing levels of salinity and nitrogen source. To address this, two glass house experiments were carried out. The first to study the shoot biomass, and nutrient accumulation response, measured by ICP-MS analysis, of edible halophyte species, including Mesembryanthemum crystallinum (ice plant), Salsola komarovii (Land seaweed), Enchylaena tomentosa (Ruby Saltbush), Crithmum maritimum (Rock Samphire), Crambe maritima (Sea Kale) and Mertensia maritima (Oyster Plant), under increasing levels of salinity (0 to 800 mM). The second experiment studied the effects of nitrogen source combined with salinity, on levels of oxalate, measured by HPLC, in ice plant and ruby saltbush. Species differences for biomass and sodium (Na), potassium (K), chloride (Cl), nitrogen (N) and phosphorus (P) accumulation were observed across the range of salt treatments (0 to 800mM). Shoot concentrations of the anti-nutrient oxalate decreased significantly in ice plant and ruby saltbush with an increase in the proportion of N provided as NH4+ (up to 100%), while shoot oxalate concentrations in ice plant and ruby saltbush grown in the absence of NaCl were not significantly different to oxalate concentrations in plants treated with 200 mM or 400 mM NaCl. However, the lower shoot oxalate concentrations observed with the increase in NH4+ came with concurrent reductions in shoot biomass. Results suggest that there will need to be a calculated tradeoff between oxalate levels and biomass when growing these plants for commercial purposes., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Farzana et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Tolerance mechanisms employed by plants under environmental stresses can protect them against other co-occurring stresses. In this study, the effect of pre-exposure and simultaneous salt treatment on nickel (Ni) toxicity tolerance in one halophyte (L. sativum) and one glycophyte (L. latifolium) Lepidium species in hydroponics was investigated. In order to compare the species independent from their salt and Ni tolerance level, the glycophyte was subjected to lower salt and Ni concentrations and for a shorter period of time than the halophyte. Salt (NaCl) was applied at 50 and 100 mM concentrations and Ni was provided at an equal free Ni 2+ activity by adding 100 and 200 µM Ni as single stresses, but 130 and 300 µM Ni for the treatment of its combination with salt in the glycophyte and halophyte, respectively. Temporal analyses of signaling molecules revealed that the halophyte is characteristically different from the glycophyte in that it exhibits a higher constitutive level of nitric oxide and hydrogen peroxide, a longer duration of response to Ni, and its augmentation by salt. In addition to higher biomass and less Ni accumulation in salt-treated plants, the concentrations of free thiol groups, leaf pigments, proline, free and cell wall-bound phenolics contents, and the activity of phenolic metabolizing enzymes were higher in L. latifolium under the combined salt and Ni treatments than under the single Ni stress. In contrast, the biomass and most biochemical parameters of Ni-stressed L. sativum plants were not enhanced by salt treatment but rather decreased. Our findings shed light on cross-tolerance mechanisms in halophytes and uncovered halophyte survival strategies under multiple stresses., (© 2022. The Author(s) under exclusive licence to The Botanical Society of Japan.)

Background: Salinization of soil is an urgent problem that restricts agroforestry production and environmental protection. Substantial accumulation of metal ions or highly alkaline soil alters plant metabolites and may even cause plant death. To explore the differences in the response strategies between Suaeda salsa (S. salsa) and Puccinellia tenuiflora (P. tenuiflora), two main constructive species that survive in saline-alkali soil, their metabolic differences were characterized., Result: Metabolomics was conducted to study the role of metabolic differences between S. salsa and P. tenuiflora under saline-alkali stress. A total of 68 significantly different metabolites were identified by GC-MS, including 9 sugars, 13 amino acids, 8 alcohols, and 34 acids. A more detailed analysis indicated that P. tenuiflora utilizes sugars more effectively and may be saline-alkali tolerant via sugar consumption, while S. salsa utilizes mainly amino acids, alcohols, and acids to resist saline-alkali stress. Measurement of phenolic compounds showed that more C6C3C6-compounds accumulated in P. tenuiflora, while more C6C1-compounds, phenolic compounds that can be used as signalling molecules to defend against stress, accumulated in S. salsa., Conclusions: Our observations suggest that S. salsa resists the toxicity of saline-alkali stress using aboveground organs and that P. tenuiflora eliminates this toxicity via roots. S. salsa has a stronger habitat transformation ability and can provide better habitat for other plants., (© 2022. The Author(s).)

Abiotic stress tolerance has been weakened during the domestication of all major staple crops. Soil salinity is a major environmental constraint that impacts over half of the world population; however, given the increasing reliance on irrigation and the lack of available freshwater, agriculture in the 21st century will increasingly become saline. Therefore, global food security is critically dependent on the ability of plant breeders to create high-yielding staple crop varieties that will incorporate salinity tolerance traits and account for future climate scenarios. Previously, we have argued that the current agricultural practices and reliance on crops that exclude salt from uptake is counterproductive and environmentally unsustainable, and thus called for a need for a major shift in a breeding paradigm to incorporate some halophytic traits that were present in wild relatives but were lost in modern crops during domestication. In this review, we provide a comprehensive physiological and molecular analysis of the key traits conferring crop halophytism, such as vacuolar Na + sequestration, ROS desensitization, succulence, metabolic photosynthetic switch, and salt deposition in trichomes, and discuss the strategies for incorporating them into elite germplasm, to address a pressing issue of boosting plant salinity tolerance., (Copyright © 2021 The Author. Published by Elsevier Inc. All rights reserved.)

Key Message: Integrative transcriptome and proteome analyses revealed many candidate members that may involve in salt secretion from salt glands in Limonium bicolor. Limonium bicolor, a typical recretohalophyte, protects itself from salt damage by excreting excess salt out of its cells through salt glands. Here, to provide an overview of the salt-tolerance mechanism of L. bicolor, we conducted integrative transcriptome and proteome analyses of this species under salt treatment. We identified numerous differentially expressed transcripts and proteins that may be related to the salt-tolerance mechanism of L. bicolor. By measuring the Na + secretion rate, were found that this cation secretion rate of a single salt gland was significantly increased after high salinity treatment compared with that in control and then reached the maximum in a short time. Interestingly, transcripts and proteins involved in transmembrane transport of ions were differentially expressed in response to high salinity treatment, suggesting a number of genes and proteins they may play important roles in the salt-stress response. Correlation between differentially expressed transcript and protein profiles revealed several transcripts and proteins that may be responsible for salt tolerance, such as cellulose synthases and annexins. Our findings uncovered many candidate transcripts and proteins in response to the salt tolerance of L. bicolor, providing deep insights into the molecular mechanisms of this important process in recretohalophytes., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Salinity is an important abiotic stress affecting plant growth. We have known that plants can recruit beneficial microbes from the surrounding soil. However, the ecological functions of the core microbiome in salt-tolerant plants, together with their driving factors, remain largely unexplored. Here, we employed both amplicon and shotgun metagenomic sequencing to investigate the microbiome and function signatures of bulk soil and rhizocompartment samples from three salt-tolerant plants (legumes Glycine soja and Sesbania cannabina and nonlegume Sorghum bicolor). Strong filtration effects for microbes and functional genes were found in the rhizocompartments following a spatial gradient. The dominant bacteria belonged to Ensifer for legumes and Bacillus for S. bicolor . Although different salt-tolerant plants harbored distinct bacterial communities, they all enriched genes involved in cell motility, Na + transport, and plant growth-promoting function (e.g., nitrogen fixation and phosphate solubilization) in rhizoplane soils, implying that the microbiome assembly of salt-tolerant plants might depend on the ecological functions of microbes rather than microbial taxa. Moreover, three metagenome-assembled genomes affiliated to Ensifer were obtained, and their genetic basis for salt stress alleviation were predicted. Soil pH, electrical conductivity, and total nitrogen were the most important driving factors for explaining the above microbial and functional gene selection. Correspondingly, the growth of an endophyte, Ensifer meliloti CL09, was enhanced by providing root exudates, suggesting that root exudates might be one of factors in the selection of rhizosphere and endosphere microbiota. Overall, this study reveals the ecological functions of the populations inhabiting the root of salt-tolerant plants. IMPORTANCE Salinity is an important but little-studied abiotic stressor affecting plant growth. Although several previous reports have examined salt-tolerant plant microbial communities, we still lack a comprehensive understanding about the functional characteristics and genomic information of this population. The results of this study revealed the root-enriched and -depleted bacterial groups, and found three salt-tolerant plants harbored different bacterial populations. The prediction of three metagenome-assembled genomes confirmed the critical role of root dominant species in helping plants tolerate salt stress. Further analysis indicated that plants enriched microbiome from soil according to their ecological functions but not microbial taxa. This highlights the importance of microbial function in enhancing plant adaptability to saline soil and implies that we should pay more attention to microbial function and not only to taxonomic information. Ultimately, these results provide insight for future agriculture using the various functions of microorganisms on the saline soil.

Background: Salt stress is one of the main constraints determining crop productivity, and therefore one of the main limitations for food production. The aim of this study was to characterize the salt stress response at the physiological and molecular level of different Broccoli (Brassica oleracea L. var. Italica Plenck) cultivars that were previously characterized in field and greenhouse trials as salt sensitive or salt tolerant. This study aimed to identify functional and molecular traits capable of predicting the ability of uncharacterized lines to cope with salt stress. For this purpose, this study measured different physiological parameters, hormones and metabolites under control and salt stress conditions., Results: This study found significant differences among cultivars for stomatal conductance, transpiration, methionine, proline, threonine, abscisic acid, jasmonic acid and indolacetic acid. Salt tolerant cultivars were shown to accumulate less sodium and potassium in leaves and have a lower sodium to potassium ratio under salt stress. Analysis of primary metabolites indicated that salt tolerant cultivars have higher concentrations of several intermediates of the Krebs cycle and the substrates of some anaplerotic reactions., Conclusions: This study has found that the energetic status of the plant, the sodium extrusion and the proline content are the limiting factors for broccoli tolerance to salt stress. Our results establish physiological and molecular traits useful as distinctive markers to predict salt tolerance in Broccoli or to design novel biotechnological or breeding strategies for improving broccoli tolerance to salt stress., (© 2021. The Author(s).)

Serine proteases are a class of versatile proteolytic enzymes. They are necessary for protein catabolism, intracellular amino acid turnover, and regulation of proteins involved in diverse molecular and cellular processes across taxa. In this study, bioinformatic analyses revealed a significantly large number of serine proteases in the halotolerant cyanobacterium Halothece sp. PCC7418 (hereafter referred to as Halothece 7418) compared to the model freshwater cyanobacterium Synechococcus elongatus PCC7942 (hereafter referred to as S. elongatus 7942). The cyanobacterial serine proteases are likely derived from different linages since no conserved motifs were detected. The presence of highly diverse serine proteases in Halothece 7418 implicated an evolutionary-mediated modification of several proteases, which may play numerous physiological roles. We also examined the gene expression patterns of 34 serine protease encoding genes in Halothece 7418 exposed to salt stress. Our results revealed that several serine protease genes were drastically up-regulated under salt with high concentration but remained unchanged under salt with low concentration. All four clp genes ( H1996, H1997, H0950 , and H3375 ) and H3553 gene (which encodes a putative HtrA protease) were significantly induced upon salt stress. These responses support the roles of the housekeeping pathways in both the degradation of damaged proteins induced by salt stress and regulation of proteins involved in the molecular recovery from salt stress. Since serine proteases share several biochemical features and physiological functions, the results from this study provide an insight into diversification of serine proteases in cyanobacteria. Further, these results will increase our understanding of several mechanisms at the subcellular level.

WRKY is one of the largest families of transcription factors in plants. It not only regulates plant growth and development but also participates in the regulation of plant defense against biological and abiotic stresses. In this study, research was aimed to overexpress WRKY39 gene of P. trichocarpa ( PtWRKY39 ) and to identify its important role played in drought and saline-alkali tolerance in tobacco model plant. Under the control of CaMV35S promoter, the overexpression of PtWRKY39 gene was increased to more than 10 times in T3 generation of transgenic tobacco plant. The drought resistance and saline-alkali tolerance were evidenced in overexpressed PtWRKY39 transgenic lines at germination/seedling stage. The overall germination rate, fresh weight, and chlorophyll contents of overexpressed lines were significantly higher while the level of malondialdehyde was significantly lower in PtWRKY39 transgenic lines than that of wild type (WT) lines. The content of H 2 O 2 in leaves was detected by the 3, 3-Diaminobenzidine method showed that the overexpression of PtWRKY39 gene could reduce the accumulation of ROS (mainly H 2 O 2 ) and enhance salt-alkali tolerance. Phenotypic analysis at 7-leaf pot transgenic seedlings stage treated with the saline-alkali soil extract and salt NaCl under root irrigation stress, revealed growth of the transgenic line was significantly higher than that of WT. This work concludes that overexpression of PtWRKY39 gene can improve the regulation of drought resistance and saline-alkali tolerance of transgenic plants during seed germination and vegetative growth.

The halophyte ice plant (Mesembryanthemum crystallinum) converts its mode of photosynthesis from C 3 to crassulacean acid metabolism (CAM) during severe water stress. During the transition to CAM, the plant induces CAM-related genes and changes its diurnal stomatal behavior to take up CO 2 efficiently at night. However, limited information concerning this signaling exists. Here, we investigated the changes in the diurnal stomatal behavior of M. crystallinum during its shift in photosynthesis using a detached epidermis. M. crystallinum plants grown under C 3 conditions opened their stomata during the day and closed them at night. However, CAM-induced plants closed their stomata during the day and opened them at night. Quantitative analysis of endogenous phytohormones revealed that trans-zeatin levels were high in CAM-induced plants. In contrast, the levels of jasmonic acid (JA) and JA-isoleucine were severely reduced in CAM-induced plants, specifically at night. CAM induction did not alter the levels of abscisic acid; however, inhibitors of abscisic acid synthesis suppressed CAM-induced stomatal closure. These results indicate that M. crystallinum regulates the diurnal balance of cytokinin and JA during CAM transition to alter stomatal behavior., (Copyright © 2021 Elsevier GmbH. All rights reserved.)

Background: Identifying genes involved in salt tolerance in the recretohalophyte Limonium bicolor could facilitate the breeding of crops with enhanced salt tolerance. Here we cloned the previously uncharacterized gene LbHLH and explored its role in salt tolerance., Results: The 2,067-bp open reading frame of LbHLH encodes a 688-amino-acid protein with a typical helix-loop-helix (HLH) domain. In situ hybridization showed that LbHLH is expressed in salt glands of L. bicolor. LbHLH localizes to the nucleus, and LbHLH is highly expressed during salt gland development and in response to NaCl treatment. To further explore its function, we heterologously expressed LbHLH in Arabidopsis thaliana under the 35S promoter. The overexpression lines showed significantly increased trichome number and reduced root hair number. LbHLH might interact with GLABRA1 to influence trichome and root hair development, as revealed by yeast two-hybrid analysis. The transgenic lines showed higher germination percentages and longer roots than the wild type under NaCl treatment. Analysis of seedlings grown on medium containing sorbitol with the same osmotic pressure as 100 mM NaCl demonstrated that overexpressing LbHLH enhanced osmotic resistance., Conclusion: These results indicate that LbHLH enhances salt tolerance by reducing root hair development and enhancing osmotic resistance under NaCl stress.

Key Message: This work demonstrates that PpCIPK1, a putative protein kinase, participates in regulating plant salt tolerance in moss Physcomitrella patens. Calcineurin B-Like protein (CBL)-interacting protein kinases (CIPKs) have been reported to be involved in multiple signaling networks and function in plant growth and stress responses, however, their biological functions in non-seed plants have not been well characterized. In this study, we report that PpCIPK1, a putative protein kinase, participates in regulating plant salt tolerance in moss Physcomitrella patens (P. patens). Phylogenetic analysis revealed that PpCIPK1 shared high similarity with its homologs in higher plants. PpCIPK1 transcription level was induced upon salt stress in P. patens. Using homologous recombination, we constructed PpCIPK1 knockout mutant lines (PpCIPK1 KO). Salt sensitivity analysis showed that independent PpCIPK1 KO plants exhibited severe growth inhibition and developmental deficiency of gametophytes under salt stress condition compared to that of wild-type P. patens (WT). Consistently, ionic homeostasis was disrupted in plants due to PpCIPK1 deletion, and high level of H 2 O 2 was accumulated in PpCIPK1 KO than that in WT. Furthermore, PpCIPK1 functions in regulating photosynthetic activity in response to salt stress. Interestingly, we observed that PpCIPK1 could completely rescue the salt-sensitive phenotype of sos2-1 to WT level in Arabidopsis, indicating that AtSOS2 and PpCIPK1 are functionally conserved. In conclusion, our work provides evidence that PpCIPK1 participates in salt tolerance regulation in P. patens.

The euhalophyte species Salicornia europaea is cultivated for oilseed and as a fodder crop in various parts of the world. In saline coastal environments it possesses great potential for the subsistence of the most disadvantaged farmers. We investigated the effect of salinity levels in irrigation water on the germination capacity, shoot biomass and seed productivity as well as diverse quality traits (nitrogen content in shoots and seeds and fatty acids, in seeds) and physiological traits (stable carbon and nitrogen isotopes and ion content) of two accessions collected in the United Arab Emirates (UAE). The three salinity levels tested were irrigation with fresh water (0.3 dS m -1 ), brackish water (25 dS m -1 ) and sea water (40 dS m -1 ). In addition, a hypersaline condition (80 dS m -1 ) was also tested for germination. The best germination rates were achieved with seeds exposed to fresh and brackish water, while imbibition with sea water decreased germination by half and hypersaline water inhibited it almost totally. However, the best irrigation regime in terms of biomass and seed yield involved brackish water. Moreover, rising salinity in the irrigation increased the stable isotope composition of carbon (δ 13 C) and nitrogen (δ 15 N), together with the Na + and K + of shoots and seeds, and the lipid levels of seeds, while the total nitrogen content and the profile of major fatty acids of seeds did not change. Differences between the two ecotypes existed for growth and seed yield with the best ecotype exhibiting lower δ 13 C and higher K + in both shoots and seeds, lower Na + and higher δ 15 N in shoots, and lower N in seeds, together with differences in major fatty acids. Physiological mechanisms behind the response to irrigation salinity and the ecotypic differences are discussed in terms of photosynthetic carbon and nitrogen metabolism., (Copyright © 2021 Elsevier B.V. All rights reserved.)

The N 6 -methyladenosine (m 6 A) modification is the most common internal post-transcriptional modification, with important regulatory effects on RNA export, splicing, stability, and translation. Studies on the m 6 A modifications in plants have focused on Arabidopsis thaliana growth and development. However, A. thaliana is a salt-sensitive and model plant species. Thus, studies aimed at characterizing the role of the m 6 A modification in the salt stress responses of highly salt-tolerant crop species are needed. Sweet sorghum is cultivated as an energy and forage crop, which is highly suitable for growth on saline-alkaline land. Exploring the m 6 A modification in sweet sorghum may be important for elucidating the salt-resistance mechanism of crops. In this study, we mapped the m 6 A modifications in two sorghum genotypes (salt-tolerant M-81E and salt-sensitive Roma) that differ regarding salt tolerance. The m 6 A modification in sweet sorghum under salt stress was drastically altered, especially in Roma, where the m 6 A modification on mRNAs of some salt-resistant related transcripts increased, resulting in enhanced mRNA stability, which in turn was involved in the regulation of salt tolerance in sweet sorghum. Although m 6 A modifications are important for regulating sweet sorghum salt tolerance, the regulatory activity is limited by the initial m 6 A modification level. Additionally, in M-81E and Roma, the differences in the m 6 A modifications were much greater than the differences in gene expression levels and are more sensitive. Our study suggests that the number and extent of m 6 A modifications on the transcripts of salt-resistance genes may be important factors for determining and assessing the salt tolerance of crops., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Saline-alkali soil has become an important environmental problem for crop productivity. One of the most effective approaches is to cultivate new stress-tolerant plants through genetic engineering. Through RNA-seq analysis and RT-PCR validation, a novel bZIP transcription factor ChbZIP1, which is significantly upregulated at alkali conditions, was obtained from alkaliphilic microalgae Chlorella sp. BLD. Overexpression of ChbZIP1 in Saccharomyces cerevisiae and Arabidopsis increased their alkali resistance, indicating ChbZIP1 may play important roles in alkali stress response. Through subcellular localization and transcriptional activation activity analyses, we found that ChbZIP1 is a nuclear-localized bZIP TF with transactivation activity to bind with the motif of G-box 2 (TGACGT). Functional analysis found that genes such as GPX1, DOX1, CAT2, and EMB, which contained G-box 2 and were associated with oxidative stress, were significantly upregulated in Arabidopsis with ChbZIP1 overexpression. The antioxidant ability was also enhanced in transgenic Arabidopsis . These results indicate that ChbZIP1 might mediate plant adaptation to alkali stress through the active oxygen detoxification pathway. Thus, ChbZIP1 may contribute to genetically improving plants' tolerance to alkali stress.

Soil salinization is a serious and growing problem around the world. Some plants, recognized as the recretohalophytes, can normally grow on saline-alkali soil without adverse effects by secreting excessive salt out of the body. The elucidation of the salt secretion process is of great significance for understanding the salt tolerance mechanism adopted by the recretohalophytes. Between the 1950s and the 1970s, three hypotheses, including the osmotic potential hypothesis, the transfer system similar to liquid flow in animals, and vesicle-mediated exocytosis, were proposed to explain the salt secretion process of plant salt glands. More recently, increasing evidence has indicated that vesicular transport plays vital roles in salt secretion of recretohalophytes. Here, we summarize recent findings, especially regarding the molecular evidence on the functional roles of vesicular trafficking in the salt secretion process of plant salt glands. A model of salt secretion in salt gland is also proposed.

Choline, as a precursor of glycine betaine (GB) and phospholipids, is known to play roles in plant tolerance to salt stress, but the downstream metabolic pathways regulated by choline conferring salt tolerance are still unclear for non-GB-accumulating species. The objectives were to examine how choline affects salt tolerance in a non-GB-accumulating grass species and to determine major metabolic pathways of choline regulating salt tolerance involving GB or lipid metabolism. Kentucky bluegrass (Poa pratensis) plants were subjected to salt stress (100 mM NaCl) with or without foliar application of choline chloride (1 mM) in a growth chamber. Choline or GB alone and the combined application increased leaf photochemical efficiency, relative water content and osmotic adjustment and reduced leaf electrolyte leakage. Choline application had no effects on the endogenous GB content and GB synthesis genes did not show responses to choline under nonstress and salt stress conditions. GB was not detected in Kentucky bluegrass leaves. Lipidomic analysis revealed an increase in the content of monogalactosyl diacylglycerol, phosphatidylcholine and phosphatidylethanolamine and a decrease in the phosphatidic acid content by choline application in plants exposed to salt stress. Choline-mediated lipid reprogramming could function as a dominant salt tolerance mechanism in non-GB-accumulating grass species., (� The Author(s) 2020. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Suaeda glauca and Puccinellia tenuiflora are two important saline-alkali tolerant plants that can improve the soil properties. For exploring the different tolerance mechanisms between them, GC-MS-based metabolomics was used to comprehensively evaluate the primary metabolites differences, a total of 51 different metabolites were present in different quantities. The identified compounds were mainly 11 sugars, 7 amino acids, 5 alcohols and 18 organic acids; they play an important role in responding to the saline-alkali stress and distinguish between S. glauca and P. tenuiflora . All identified metabolites classes showed similar trend to largely accumulate in P. tenuiflora roots and S. glauca shoots, this reveals that the two plants used different physiological strategies to cope with saline-alkali stress.

Arabidopsis thaliana TRY is a negative regulator of trichome differentiation that promotes root hair differentiation. Here, we established that LbTRY, from the recretohalophyte Limonium bicolor, is a typical MYB transcription factor that exhibits transcriptional activation activity and locates in nucleus. By in situ hybridization in L. bicolor, LbTRY may be specifically positioned in salt gland of the expanded leaves. LbTRY expression was the highest in mature leaves and lowest under NaCl treatment. For functional assessment, we heterologously expressed LbTRY in wild-type and try29760 mutant Arabidopsis plants. Epidermal differentiation was remarkably affected in the transgenic wild-type line, as was increased root hair development. Complementation of try29760 with LbTRY under both 35S and LbTRY specific promoter restored the wild-type phenotype. qRT-PCR analysis suggested that AtGL3 and AtZFP5 promote root hair cell fate in lines heterologously producing LbTRY. In addition, four genes (AtRHD6, AtRSL1, AtLRL2, and AtLRL3) involved in root hair initiation and elongation were upregulated in the transgenic lines. Furthermore, LbTRY specifically increased the salt sensitivity of the transgenic lines. The transgenic and complementation lines showed poor germination rates and reduced root lengths, whereas the mutant unexpectedly fared the best under a range of NaCl treatments. Under salt stress, the transgenic seedlings accumulated more MDA and Na + and less proline and soluble sugar than try29760. Thus, when heterologously expressed in Arabidopsis, LbTRY participates in hair development, similar to other MYB proteins, and specifically reduces salt tolerance by increasing ion accumulation and reducing osmolytes. The expression of salt-tolerance marker genes (SOS1, SOS2, SOS3 and P5CS1) was significant reduced in the transgenic lines. More will be carried by downregulating expression of TRY homologs in crops to improve salt tolerance., (Copyright © 2020 The Authors. Published by Elsevier B.V. All rights reserved.)

Backgrounds: Pomegranate (Punica granatum L.) is an important commercial fruit tree, with moderate tolerance to salinity. The balance of Cl - and other anions in pomegranate tissues are affected by salinity, however, the accumulation patterns of anions are poorly understood. The chloride channel (CLC) gene family is involved in conducting Cl - , NO 3 - , HCO 3 - and I - , but its characteristics have not been reported on pomegranate., Results: In this study, we identified seven PgCLC genes, consisting of four antiporters and three channels, based on the presence of the gating glutamate (E) and the proton glutamate (E). Phylogenetic analysis revealed that seven PgCLCs were divided into two clades, with clade I containing the typical conserved regions GxGIPE (I), GKxGPxxH (II) and PxxGxLF (III), whereas clade II not. Multiple sequence alignment revealed that PgCLC-B had a P [proline, Pro] residue in region I, which was suspected to be a NO 3 - /H + exchanger, while PgCLC-C1, PgCLC-C2, PgCLC-D and PgCLC-G contained a S [serine, Ser] residue, with a high affinity to Cl - . We determined the content of Cl - , NO 3 - , H 2 PO 4 - , and SO 4 2- in pomegranate tissues after 18 days of salt treatments (0, 100, 200 and 300 mM NaCl). Compared with control, the Cl - content increased sharply in pomegranate tissues. Salinity inhibited the uptake of NO 3 - and SO 4 2- , but accelerated H 2 PO 4 - uptake. The results of real-time reverse transcription PCR (qRT-PCR) revealed that PgCLC genes had tissue-specific expression patterns. The high expression levels of three antiporters PgCLC-C1, PgCLC-C2 and PgCLC-D in leaves might be contributed to sequestrating Cl - into the vacuoles. However, the low expression levels of PgCLCs in roots might be associated with the exclusion of Cl - from root cells. Also, the up-regulated PgCLC-B in leaves indicated that more NO 3 - was transported into leaves to mitigate the nitrogen deficiency., Conclusions: Our findings suggested that the PgCLC genes played important roles in balancing of Cl - and NO 3 - in pomegranate tissues under salt stress. This study established a theoretical foundation for the further functional characterization of the CLC genes in pomegranate.

Soil salinization is a major threat to global food security and the biodiversity of natural ecosystems. To adapt to salt stress, plants rely on ROS-mediated signalling networks that operate upstream of a broad array of physiological and genetic processes. A key player in ROS signalling is NADPH oxidase, a plasma-membrane-bound enzyme encoded by RBOH genes. In this study, we have conducted a comprehensive bioinformatic analysis of over 50 halophytic and glycophytic species to link the difference in the kinetics of ROS signalling between contrasting species with the abundance and/or structure of NADPH oxidases. The RBOH proteins were predicted in all the tested plant lineages except some algae species from the Rhodophyta, Chlorophyta and Streptophyta. Within the glycophytic group, the number of RBOH copies correlated negatively with salinity stress tolerance, suggesting that a reduction in the number of RBOH isoforms may be potentially related to the evolution of plant salinity tolerance. While halophytes did not develop unique protein families during evolution, they evolved additional phosphorylation target sites at the N-termini of NADPH oxidases, potentially modulating enzyme activity and allowing more control over their function, resulting in more efficient ROS signalling and adaptation to saline conditions., (© 2020 John Wiley & Sons Ltd.)

Under certain abiotic conditions (elevated irradiance, temperature and sediment salinity) observed mostly during the Mediterranean summer, the halophyte Sarcocornia fruticosa suffers a metabolic shift evidenced by a red coloration, evidencing the presence of two physiotypes (green and red). Previous works indicated that this metabolic shift has severe implications in the primary photochemistry of this species, impairing the light and carbon harvesting. Under stress plants have lower light use efficiencies and are more prone to photoinhibition, and thus this metabolic shift is essential for this species to deal with the high light intensities characteristic from this time of the year. Nevertheless, the fatty acid and lipid remodelling in green and red S. fruticosa physiotypes was not previously evaluated nor its relations with this metabolic shift. The evaluation of the lipid landscape suggests several lipid and fatty acid remodelling when comparing both red and green physiotype, as strategies to overcome stress. The galactolipids of the red physiotype suffer several changes aiming to keep chloroplast membrane structural and functional stability during water stress and can also be related to an improvement of the plants response to osmotic stress. At the phospholipid level, a readjustment of its fatty acid profiles was also observable. This remodelling allows the plants to adjust membrane fluidity the imposed osmotic stress, being this action transversal to choroplastidial, extraplastidial, and involves the action of the different phospholipids. Additionally, neutral lipids (NLs) also appear to play a role in osmotic stress adaptation, with an increase content in C18 fatty acids in the red physiotype. The resulting lipid landscape in both physiotypes presents very specific signatures that can be used as biomarkers to track this kind of metabolic shifts, in future studies with similar species., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Salinization of land is likely to increase due to climate change with impact on agricultural production. Since most species used as crops are sensitive to salinity, improvement of salt tolerance is needed to maintain global food production. This review summarises successes and failures of transgenic approaches in improving salt tolerance in crop species. A conceptual model of coordinated physiological mechanisms in roots and shoots required for salt tolerance is presented. Transgenic plants overexpressing genes of key proteins contributing to Na + 'exclusion' (PM-ATPases with SOS1 antiporter, and HKT1 transporter) and Na + compartmentation in vacuoles (V-H + ATPase and V-H + PPase with NHX antiporter), as well as two proteins potentially involved in alleviating water deficit during salt stress (aquaporins and dehydrins), were evaluated. Of the 51 transformations, with gene(s) involved in Na + 'exclusion' or Na + vacuolar compartmentation that contained quantitative data on growth and include a non-saline control, 48 showed improvements in salt tolerance (less impact on plant mass) of transgenic plants, but with only two tested in field conditions. Of these 51 transformations, 26 involved crop species. Tissue ion concentrations were altered, but not always in the same way. Although glasshouse data are promising, field studies are required to assess crop salinity tolerance., (© 2020 John Wiley & Sons Ltd.)

The Plumbaginaceae (non-core Caryophyllales) is a family well known for species adapted to a wide range of arid and saline habitats. Of its salt-tolerant species, at least 45 are in the genus Limonium; two in each of Aegialitis, Limoniastrum and Myriolimon, and one each in Psylliostachys, Armeria, Ceratostigma, Goniolimon and Plumbago. All the halophytic members of the family have salt glands and salt glands are also common in the closely related Tamaricaceae and Frankeniaceae. The halophytic species of the three families can secrete a range of ions (Na + , K + , Ca 2+ , Mg 2+ , Cl - , HCO 3 - , SO 4 2- ) and other elements (As, Cd, Cr, Cu, Fe, Mn, Ni, Pb and Zn). Salt glands are, however, absent in salt-tolerant members of the sister family Polygonaceae. We describe the structure of the salt glands in the three families and consider whether glands might have arisen as a means to avoid the toxicity of Na + and/or Cl - or to regulate Ca 2+ concentrations with the leaves. We conclude that the establishment of lineages with salt glands took place after the split between the Polygonaceae and its sister group the Plumbaginaceae., (© 2020 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Key Message: PvArf regulate proline biosynthesis by physically interacting with PvP5CS1 to improve salt tolerance in switchgrass. The genetic factors that contribute to stress resiliency are yet to be determined. Here, we identified three ADP-ribosylation factors, PvArf1, PvArf-B1C, and PvArf-related, which contribute to salinity tolerance in transgenic switchgrass (Panicum virgatum L.). Switchgrass overexpressing each of these genes produced approximately twofold more biomass than wild type (WT) under normal growth conditions. Transgenic plants accumulated modestly higher levels of proline under normal conditions, but this level was significantly increased under salt stress providing better protection to transgenic plants compared to WT. We found that PvArf genes induce proline biosynthesis genes under salt stress to positively regulate proline accumulation, and further demonstrated that PvArf physically interact with PvP5CS1. Moreover, the transcript levels of two key ROS-scavenging enzyme genes were significantly increased in the transgenic plants compared to WT, leading to reduced H 2 O 2 accumulation under salt stress conditions. PvArf genes also protect cells against stress-induced changes in Na + and K + ion concentrations. Our findings uncover that ADP-ribosylation factors are key determinants of biomass yield in switchgrass, and play pivotal roles in salinity tolerance by regulating genes involved in proline biosynthesis.

The adaptation strategies of halophytic seaside barley Hordeum marinum to high salinity and osmotic stress were investigated by nuclear magnetic resonance imaging, as well as ionomic, metabolomic, and transcriptomic approaches. When compared with cultivated barley, seaside barley exhibited a better plant growth rate, higher relative plant water content, lower osmotic pressure, and sustained photosynthetic activity under high salinity, but not under osmotic stress. As seaside barley is capable of controlling Na + and Cl - concentrations in leaves at high salinity, the roots appear to play the central role in salinity adaptation, ensured by the development of thinner and likely lignified roots, as well as fine-tuning of membrane transport for effective management of restriction of ion entry and sequestration, accumulation of osmolytes, and minimization of energy costs. By contrast, more resources and energy are required to overcome the consequences of osmotic stress, particularly the severity of reactive oxygen species production and nutritional disbalance which affect plant growth. Our results have identified specific mechanisms for adaptation to salinity in seaside barley which differ from those activated in response to osmotic stress. Increased knowledge around salt tolerance in halophytic wild relatives will provide a basis for improved breeding of salt-tolerant crops.

Background: Chitosan (CTS), a natural polysaccharide, exhibits multiple functions of stress adaptation regulation in plants. However, effects and mechanism of CTS on alleviating salt stress damage are still not fully understood. Objectives of this study were to investigate the function of CTS on improving salt tolerance associated with metabolic balance, polyamine (PAs) accumulation, and Na + transport in creeping bentgrass (Agrostis stolonifera)., Results: CTS pretreatment significantly alleviated declines in relative water content, photosynthesis, photochemical efficiency, and water use efficiency in leaves under salt stress. Exogenous CTS increased endogenous PAs accumulation, antioxidant enzyme (SOD, POD, and CAT) activities, and sucrose accumulation and metabolism through the activation of sucrose synthase and pyruvate kinase activities, and inhibition of invertase activity. The CTS also improved total amino acids, glutamic acid, and γ-aminobutyric acid (GABA) accumulation. In addition, CTS-pretreated plants exhibited significantly higher Na + content in roots and lower Na + accumulation in leaves then untreated plants in response to salt stress. However, CTS had no significant effects on K + /Na + ratio. Importantly, CTS enhanced salt overly sensitive (SOS) pathways and also up-regulated the expression of AsHKT1 and genes (AsNHX4, AsNHX5, and AsNHX6) encoding Na + /H + exchangers under salt stress., Conclusions: The application of CTS increased antioxidant enzyme activities, thereby reducing oxidative damage to roots and leaves. CTS-induced increases in sucrose and GABA accumulation and metabolism played important roles in osmotic adjustment and energy metabolism during salt stress. The CTS also enhanced SOS pathway associated with Na + excretion from cytosol into rhizosphere, increased AsHKT1 expression inhibiting Na + transport to the photosynthetic tissues, and also up-regulated the expression of AsNHX4, AsNHX5, and AsNHX6 promoting the capacity of Na + compartmentalization in roots and leaves under salt stress. In addition, CTS-induced PAs accumulation could be an important regulatory mechanism contributing to enhanced salt tolerance. These findings reveal new functions of CTS on regulating Na + transport, enhancing sugars and amino acids metabolism for osmotic adjustment and energy supply, and increasing PAs accumulation when creeping bentgrass responds to salt stress.

This study aimed to reveal the mechanistic basis of the melatonin-mediated amelioration of salinity stress in plants. Electrophysiological experiments revealed that melatonin decreased salt-induced K + efflux (a critical determinant of plant salt tolerance) in a dose- and time-dependent manner and reduced sensitivity of the plasma membrane K + -permeable channels to hydroxyl radicals. These beneficial effects of melatonin were abolished by NADPH oxidase blocker DPI. Transcriptome analyses revealed that melatonin induced 585 (448 up- and 137 down-regulated) and 59 (54 up- and 5 down-regulated) differentially expressed genes (DEGs) in the root tip and mature zone, respectively. The most noticeable changes in the root tip were melatonin-induced increase in the expression of several DEGs encoding respiratory burst NADPH oxidases (OsRBOHA and OsRBOHF), calcineurin B-like/calcineurin B-like-interacting protein kinase (OsCBL/OsCIPK), and calcium-dependent protein kinase (OsCDPK) under salt stress. Melatonin also enhanced the expression of potassium transporter genes (OsAKT1, OsHAK1, and OsHAK5). Taken together, these results indicate that melatonin improves salt tolerance in rice by enabling K + retention in roots, and that the latter process is conferred by melatonin scavenging of hydroxyl radicals and a concurrent OsRBOHF-dependent ROS signalling required to activate stress-responsive genes and increase the expression of K + uptake transporters in the root tip., (© 2020 John Wiley & Sons Ltd.)

Plant salt tolerance is a complex mechanism, and different plant species have different strategies for surviving salt stress. In the present study, we analyzed and compared the morphological and physiological responses of two willow species (Salix linearistipularis and Salix matsudana) from different habitats to salt stress. S. linearistipularis exhibited higher seed germination rates and seedling root Na + efflux than S. matsudana under salt stress. After salt treatment, S. linearistipularis leaves exhibited less Na + accumulation, loss of water and chlorophyll, reduction in photosynthetic capacity, and damage to leaf cell structure than leaves of S. matsudana. Scanning electron microscopy combined with gas chromatography mass spectrometry showed that S. linearistipularis leaves had higher cuticular wax loads than S. matsudana leaves. Overall, our results showed that S. linearistipularis had higher salt tolerance than S. matsudana, which was associated with different morphological and physiological responses to salt stress. Furthermore, our study suggested that S. linearistipularis could be a promising tree species for saline-alkali land greening and improvement.

Background: This study describes a promising method for understanding how halophytes adapt to extreme saline conditions and to identify populations with greater resistance. Image and colour analyses have the ability to obtain many image parameters and to discriminate between different aspects in plants, which makes them a suitable tool in combination with genetic analysis to study the plants salt tolerance. To the best of our knowledge, there are no publications about the monitoring of halophytic plants by non-destructive methods for identifying the differences between plants that belong to different maternal salinity environments. The aim is to evaluate the ability of image analysis as a non-destructive method and principal component analysis (PCA) to identify the multiple responses of two S. europaea populations, and to determine which population is most affected by different salinity treatments as a preliminary model of selection., Results: Image analysis was beneficial for detecting the phenotypic variability of two S. europaea populations by morphometric and colour parameters, fractal dimension (FD), projected area (A), shoot height (H), number of branches (B), shoot diameter (S) and colour change (ΔE). S was found to strongly positively correlate with both proline content and ΔE, and negatively with chlorophyll content. These results suggest that proline and ΔE are strongly linked to plant succulence, while chlorophyll decreases with increased succulence. The negative correlation between FD and hydrogen peroxide (HP) suggests that when the plant is under salt stress, HP content increases in plants causing a reduction in plant complexity and foliage growth. The PCA results indicate that the greater the stress, the more marked the differences. At 400 mM a shorter distance between the factorial scores was observed. Genetic variability analysis provided evidence of the differences between these populations., Conclusions: Our non-destructive method is beneficial for evaluating the halophyte development under salt stress. FD, S and ΔE were relevant indicators of plant architecture. PCA provided evidence that anthropogenic saline plants were more tolerant to saline stress. Furthermore, random amplified polymorphic DNA analysis provided a quick method for determining genetic variation patterns between the two populations and provided evidence of genetic differences between them.

Vesicular trichomes play a key role in excluding toxic ions from some halophyte species, preventing the essential processes and functions of plants from being altered. Thus, the present study aimed to evaluate the influence of these structures on Atriplex nummularia irrigated using waters with three levels of osmotic potential (-0.1, -1.4 and -2.7 MPa), formulated with NaCl in plants with vesicular trichomes and plants with partial removal of trichomes. The experiment was conducted in a protected environment and plants were evaluated for physiological parameters (water, osmotic and pressure potentials, relative water content, osmotic adjustment, pressure-volume curve, gas exchange), electrolyte leakage, lipid peroxidation and enzymatic activity (superoxide dismutase, ascorbate peroxidase, catalase). The results obtained made it possible to identify the strong contribution of vesicular trichomes to physiological and biochemical parameters, with indication of cell wall stiffening and maintenance of turgor. Furthermore, the evaluation of the osmotic potentials obtained in the study suggests that the contribution of vesicular trichomes to the salinity tolerance of the species is greater than that of osmotic adjustment. Furthermore, gas exchange results suggest that the presence of trichomes was able to regulate stomatal processes so that the plant maintains its photosynthetic performance. Evaluation of electrolyte leakage, together with the increase in malondialdehyde content, showed that the maintenance of trichomes reduces the probability of oxidative stress. The activity of antioxidant enzymes was efficient in eliminating reactive oxygen species, especially the activity of ascorbate peroxidase, which stood out in terms of hydrogen peroxide detoxification., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Background: Bread wheat is one of the most important crops for the human diet, but the increasing soil salinization is causing yield reductions worldwide. Improving salt stress tolerance in wheat requires the elucidation of the mechanistic basis of plant response to this abiotic stress factor. Although several studies have been performed to analyze wheat adaptation to salt stress, there are still some gaps to fully understand the molecular mechanisms from initial signal perception to the onset of responsive tolerance pathways. The main objective of this study is to exploit the dynamic salt stress transcriptome in underlying QTL regions to uncover candidate genes controlling salt stress tolerance in bread wheat. The massive analysis of 3'-ends sequencing protocol was used to analyze leave samples at osmotic and ionic phases. Afterward, stress-responsive genes overlapping QTL for salt stress-related traits in two mapping populations were identified., Results: Among the over-represented salt-responsive gene categories, the early up-regulation of calcium-binding and cell wall synthesis genes found in the tolerant genotype are presumably strategies to cope with the salt-related osmotic stress. On the other hand, the down-regulation of photosynthesis-related and calcium-binding genes, and the increased oxidative stress response in the susceptible genotype are linked with the greater photosynthesis inhibition at the osmotic phase. The specific up-regulation of some ABC transporters and Na + /Ca 2+ exchangers in the tolerant genotype at the ionic stage indicates their involvement in mechanisms of sodium exclusion and homeostasis. Moreover, genes related to protein synthesis and breakdown were identified at both stress phases. Based on the linkage disequilibrium blocks, salt-responsive genes within QTL intervals were identified as potential components operating in pathways leading to salt stress tolerance. Furthermore, this study conferred evidence of novel regions with transcription in bread wheat., Conclusion: The dynamic transcriptome analysis allowed the comparison of osmotic and ionic phases of the salt stress response and gave insights into key molecular mechanisms involved in the salt stress adaptation of contrasting bread wheat genotypes. The leveraging of the highly contiguous chromosome-level reference genome sequence assembly facilitated the QTL dissection by targeting novel candidate genes for salt tolerance.

In this study, we characterized, identified, and determined the effect of salt-tolerant PGPR isolated from coastal saline areas on rice growth and yield. A total of 44 bacterial strains were isolated, and 5 were found to be tolerant at high salt concentration. These isolates were further characterized for salinity tolerance and beneficial traits through a series of quantitative tests. Biochemical characterization showed that bacterial survivability decreases gradually with the increase of salt concentration. One of the strains, UPMRB9, produced the highest amount of exopolysaccharides when exposed to 1.5M of NaCl. Moreover, UPMRB9 absorbed the highest amount of sodium from the 1.5M of NaCl-amended media. The highest floc yield and biofilm were produced by UPMRE6 and UPMRB9 respectively, at 1M of NaCl concentration. The SEM observation confirmed the EPS production of UPMRB9 and UPMRE6 at 1.5M of NaCl concentration. These two isolates were identified as Bacillus tequilensis and Bacillus aryabhattai based on the 16S rRNA gene sequence. The functional group characterization of EPS showed the presence of hydroxyl, carboxyl, and amino groups. This corresponded to the presence of carbohydrates and proteins in the EPS and glucose was identified as the major type of carbohydrate. The functional groups of EPS can help to bind and chelate Na+ in the soil and thereby reduces the plant's exposure to the ion under saline conditions. The plant inoculation study revealed significant beneficial effects of bacterial inoculation on photosynthesis, transpiration, and stomatal conductance of the plant which leads to a higher yield. The Bacillus tequilensis and Bacillus aryabhattai strains showed good potential as PGPR for salinity mitigation practice for coastal rice cultivation., Competing Interests: The authors have declared that no competing interests exist