Synergistic effects of carbon cycle metabolism and photosynthesis in Chinese cabbage under salt stress

Chinese cabbage(Brassica rapa ssp. pekinensis) has a long cultivation history and is one of the vegetable crops with the largest cultivation area in China. However, salt stress severely damages photosynthesis and hormone metabolism, nutritional balances, and results in ion toxicity in plants. To better understand the mechanisms of salt-induced growth inhibition in Chinese cabbage, RNA-seq and physiological index determination were conducted to explore the impacts of salt stress on carbon cycle metabolism and photosynthesis in Chinese cabbage. Here, we found that the number of thylakoids and grana lamellae and the content of starch granules and chlorophyll in the leaves of Chinese cabbage under salt stress showed a time-dependent response, first increasing and then decreasing. Chinese cabbage increased the transcript levels of genes related to the photosynthetic apparatus and carbon metabolism under salt stress, probably in an attempt to alleviate damage to the photosynthetic system and enhance CO_(2) fixation and energy metabolism. The transcription of genes related to starch and sucrose synthesis and degradation were also enhanced;this might have been an attempt to maintain intracellular osmotic pressure by increasing soluble sugar concentrations. Soluble sugars could also be used as potential reactive oxygen species(ROS) scavengers, in concert with peroxidase(POD)enzymes, to eliminate ROS that accumulate during metabolic processes. Our study characterizes the synergistic response network of carbon metabolism and photosynthesis under salt stress.

Melatonin Alleviates Abscisic Acid Deficiency Inhibition on Photosynthesis and Antioxidant Systems in Rice under Salt Stress

Melatonin and abscisic acid,as major plant hormones,play important roles in the physiological and biochemical activities of crops,but the interaction between the two under salt stress is not yet clear.This study investigated the endogenous levels of melatonin and abscisic acid in rice by using exogenous melatonin,abscisic acid,and their synthetic inhibitors,and examined their interactions under salt stress.The research results indicate that melatonin and abscisic acid can improve rice salt tolerance.Melatonin alleviated the salt sensitivity caused by abscisic acid deficiency,increased antioxidant enzyme activity and antioxidant content in rice treated with abscisic acid synth-esis inhibitors,and reduced total reactive oxygen species content and thiobarbituric acid reactive substance accu-mulation.Melatonin also increased the activity of key photosynthetic enzymes and the content of photosynthetic pigments,maintaining the parameters of photosynthetic gas exchange and chlorophyllfluorescence.In summary,melatonin alleviated the effects of abscisic acid deficiency on photosynthesis and antioxidant systems in rice and improved salt tolerance.This study is beneficial for expanding the understanding of melatonin regulation of crop salt tolerance.

EuSHT Acts as a Hub Gene Involved in the Biosynthesis of 6-Hydroxyluteolin and Quercetin Induced by Salt Stress in Eucommia ulmoides

Salt stress inhibits plant growth and affects the biosynthesis of its secondary metabolites.Flavonoids are natural compounds that possess many important biological activities,playing a significant role in the medicinal activity of Eucommia ulmoides(E.ulmoides).To investigate the mechanism by which salt stress affects the biosynthesis of flavonoids in E.ulmoides,a comprehensive analysis of metabolomics and transcriptomics was conducted.The results indicated that salt stress led to the wilting and darkening of E.ulmoides leaves,accompanied by a decrease in chlorophyll levels,and significantly induced malondialdehyde(MDA)and relative electrical conductivity.During salt stress,most metabolites in the flavonoid biosynthesis pathway of E.ulmoides were upregulated,indicating that flavonoid biosynthesis is likely the main induced pathway under salt stress.Among them,secondary metabolites such as 6-Hydroxyluteolin and Quercetin are likely key metabolites induced by salt stress.The correlation analysis of transcriptomics and metabolomics revealed that EuSHT is a hub gene induced by salt stress,promoting the production of flavonoids such as 6-Hydroxyluteolin.The co-expression network showed a strong positive correlation between EuSHT and the biosynthesis of 6-Hydroxyluteolin and Quercetin,while it exhibited a negative correlation with Catechin biosynthesis.The branches leading to Luteolin and Dihydroquercetin are likely the main pathways for flavonoid compound biosynthesis in the plant stress response during salt stress.The results of this study provided a preliminary mechanism of secondary metabolites such as flavonoids in the medicinal plant E.ulmoides induced by salt stress and provided new theoretical support for discussing the mechanism of plant stress response.It also provided useful information for subsequent exploration of resistance genes in E.ulmoides.

Genome-Wide Exploration of the Grape GLR Gene Family and Differential Responses of VvGLR3.1 and VvGLR3.2 to Low Temperature and Salt Stress

Grapes,one of the oldest tree species globally,are rich in vitamins.However,environmental conditions such as low temperature and soil salinization significantly affect grape yield and quality.The glutamate receptor(GLR)family,comprising highly conserved ligand-gated ion channels,regulates plant growth and development in response to stress.In this study,11 members of the VvGLR gene family in grapes were identified using whole-genome sequence analysis.Bioinformatic methods were employed to analyze the basic physical and chemical properties,phylogenetic trees,conserved domains,motifs,expression patterns,and evolutionary relationships.Phylogenetic and collinear analyses revealed that the VvGLRs were divided into three subgroups,showing the high conservation of the grape GLR family.These members exhibited 2 glutamate receptor binding regions(GABAb and GluR)and 3-4 transmembrane regions(M1,M2,M3,and M4).Real-time quantitative PCR analysis demonstrated the sensitivity of all VvGLRs to low temperature and salt stress.Subsequent localization studies in Nicotiana tabacum verified that VvGLR3.1 and VvGLR3.2 proteins were located on the cell membrane and cell nucleus.Additionally,yeast transformation experiments confirmed the functionality of VvGLR3.1 and VvGLR3.2 in response to low temperature and salt stress.Thesefindings highlight the significant role of the GLR family,a highly conserved group of ion channels,in enhancing grape stress resistance.This study offers new insights into the grape GLR gene family,providing fundamental knowledge for further functional analysis and breeding of stress-resistant grapevines.

Different Approaches to Reduce Salinity in Salt-Affected Soils and Enhancing Salt Stress Tolerance in Plants

Salt stress is one of the most harmful environmental stresses in recent times and represents a significant threat to food security. Soil salinization is caused by spontaneous natural processes of mineral dissolution and human activities such as inappropriate irrigation practices. Natural geological progressions like weathering of rocks, arid climate, and higher evaporation, as well as anthropogenic activities, including the use of brackish water for irrigation, and poor tillage operations, are the foremost causes of soil salinization. Typical characteristics of saline soils are salt stress, high pH, and lack of organic carbon, as well as low availability of nutrients. Disruption of precipitation patterns as well as high average annual temperatures due to climate change additionally negatively affects the process of soil salinization. Productivity and ability to support crop growth are reduced on saline soil. Salinity-induced stress reduces plant growth by modulating the antioxidative system and nutrient orchestration. The aim of this work is to show that the mentioned problems can be alleviated in several ways such as the addition of biochar, exogenous application of several elicitors, seed priming, etc. Research has shown that the addition of biochar can significantly improve the recovery of saline soil. The addition of biochar has no significant effect on soil pH, while the cation exchange capacity of the soil increased by 17%, and the electrical conductivity of the saturated paste extract decreased by 13.2% (depends on the initial salinity and the type of biochar raw material). Moreover, biochar enriched with silicon increases the resistance of bananas to salt stress. In addition, exogenous application of several elicitors helps plants to alleviate stress by inducing stress-related physicochemical and molecular changes (selenium, sulfur, silicon, salicylic acid). Finally, seed priming showed positive effects on metabolomics, proteomics and growth of plants subjected to abiotic stress. Priming usually involves immersing the seed in a solution for a period of time to induce physiological and metabolic progression prior to germination.

Effects of Different Arbuscular Mycorrhizal Fungi on Physiology of Viola prionantha under Salt Stress

Arbuscular mycorrhizal(AM)fungi distribute widely in natural habits and play a variety of ecological functions.In order to test the physiological response to salt stress mediated by different AM fungi,Viola prionantha was selected as the host,the dominant AM fungus in the rhizosphere of V.philippica growing in Songnen saline-alkali grassland,Rhizophagus irregularis,and their mixtures were used as inoculants,and NaCl stress was applied after the roots were colonized.The results showed that V.philippica could be colonized by AM fungi in the field and the colonization rate ranged from 73.33%to 96.67%,and Claroideoglomus etunicatum was identified as the dominant AM fungi species in the rhizosphere of V.philippica by morphology combined with sequencing for AM fungal AML1/AML2 target.Inoculation with both the species resulted in the formation of mycorrhizal symbiosis(the colonization rate was more than 70%)and AM fungi significantly enhanced plants’tolerance to salt stress of varying magnitude.Higher activity of antioxidant enzymes and augmented levels of proline and other osmoregulators were observed in AM plants.The content of MDA in CK was higher than that in the inoculations with the stress of 100,200,and 250 mM.All indices except soluble protein content and MDA content were significantly correlated with AM fungal colonization indices.The analysis for different AM fungal effects showed that the mixtures and R.irregularis worked even better than C.etunicatum.These results will provide theoretical support for the exploration and screening of salt-tolerant AM fungi species and also for the application of AM-ornamental plants in saline-alkali urban greening.

The lectin gene TRpL1 of tetraploid Robinia pseudoacacia L.response to salt stress

Lectins are natural proteins in animals,plants,and microorganisms and can be divided into 12 families.These lectins play important roles in various environmental stresses.Some polyploid plants show tolerance to environmental stresses and to insect pests.However,the mechanism of stress tolerance is unclear.Tetraploid Robinia pseudoacacia(4×)under salt stress showed higher tolerance than diploid R.pseudoacacia(2×).As lectin can improve stress tolerance,it was questioned whether the stress resistance of polyploid plants was related to the lectin protein.In this study,salt resistance of lectin gene TRpL1 was verified by its over-expression in plants.In addition,salt resistance of lectin protein by E.coli strains was detected.The data revealed that the over-expression transgenic plants of TRpL1 showed better salt tolerance than control plants under salt stress,and the TRpL1-expressing strain also grew better in the medium with added NaCl.Therefore,tetraploid plants can resist salt stress through TRpL1 protein regulation.

Identification of novel salt stress-responsive microRNAs through sequencing and bioinformatic analysis in a unique halophilic Dunaliella salina strain

Dunaliella salina is a classic halophilic alga.However,its molecular mechanisms in response to high salinity at the post transcriptional level remain unknown.A unique halophilic alga strain,DS-CN1,was screened from four D.salina strains via cell biological,physiological,and biochemical methods.High-throughput sequencing of small RNAs(sRNAs)of DS-CN1 in culture medium containing 3.42-mol/L NaCl(SS group)or 0.05-mol/L NaCl(CO group)was performed on the BGISEQ-500 platform.The annotation and sequences of D.salina sRNAs were profiled.Altogether,44 novel salt stress-responsive microRNAs(miRNAs)with a relatively high C content,with the majority of them being 24 nt in length,were identified and characterized in DS-CN1.Twenty-one differentially expressed miRNAs(DEMs)in SS and CO were screened via bioinformatic analysis.A total of 319 putative salt stress-related genes targeted(104 overlapping genes)by novel miRNAs in this alga were screened based on our previous transcriptome sequencing research.Furthermore,these target genes were classified and enriched by GO and KEGG pathway analysis.Moreover,5 novel DEMs(dsa-mir3,dsa-mir16,dsa-mir17,and dsa-mir26 were significantly upregulated,and dsa-mir40 was significantly downregulated)and their corresponding 10 target genes involved in the 6 significantly enriched metabolic pathways were verified by quantitative real-time PCR.Next,their regulatory relationships were comprehensively analyzed.Lastly,a unique salt stress response metabolic network was constructed based on the novel DEM-target gene pairs.Taken together,our results suggest that 44 novel salt stress-responsive microRNAs were identified,and 4 of them might play important roles in D.salina upon salinity stress and contribute to clarify its distinctive halophilic feature.Our study will shed light on the regulatory mechanisms of salt stress responses.

GhSCL4 Acts as a Positive Regulator in Both Transgenic Arabidopsis and Cotton during Salt Stress

GRAS transcription factors play important roles in plant abiotic stress response,but their characteristics and functions in cotton have not been fully investigated.A cotton SCL4/7 subgroup gene in the GRAS family,GhSCL4,was found to be induced by NaCl treatments.Nuclear localization and transactivation activity of GhSCL4 indicate its potential role in transcriptional regulation.Transgenic Arabidopsis thaliana over-expressing GhSCL4 showed enhanced resistance to salt and osmotic stress.What’s more,the transcript levels of salt stress-induced genes(AtNHX1 and AtSOS1)and oxidation-related genes(AtAPX3 and AtCSD2)were more highly induced in the GhSCL4 over-expression lines than in wild type after salt treatment.Furthermore,silencing of GhSCL4 resulted in reduced salt tolerance in cotton caused by reactive oxygen species(ROS)enrichment under salt treatment,and antioxidant enzyme activities were accordingly significantly reduced in GhSLC4-silenced lines.These results indicated that GhSCL4 enhances salt tolerance of cotton by detoxifying ROS.In addition,the transient expression assay confirmed an interactive relationship between GhSCL4 and GhCaM7,which indicated that salt tolerance conferred by GhSCL4 might be associated with salt-induced Ca^(2+)/CaM7-mediated signaling.Taken together,GhSCL4 acts as a positive regulator in cotton during salt stress that is potentially useful for engineering salt-tolerant cotton.

Proteome-wide identification of S-sulfenylated cysteines response to salt stress in Brassica napus root

Reactive oxygen species(ROS)play a key role in a variety of biological processes,such as the perception of abiotic stress,the integration of different environmental signals,and the activation of stress response networks.Salt stress could induce an increased ROS accumulation in plants,disrupting intracellular redox homeostasis,leading to posttranslational modifications(PTMs)of specific proteins,and eventually causing adaptive changes in metabolism.Here,we performed an iodoTMT-based proteomic approach to identify the sulfenylated proteins in B.napus root responsing to salt stress.Totally,1348 sulfenylated sites in 751 proteins were identified and these proteins were widely existed in different cell compartments and processes.Our study revealed that proteins with changed abundance and sulfenylation level in B.napus root under salt stress were mainly enriched in the biological processes of ion binding,glycolysis,ATP binding,and oxidative stress response.This study displays a landscape of sulfenylated proteins response to salt stress in B.napus root and provides some theoretical support for further understanding of the molecular mechanisms of redox regulation under salt stress in plants.

Chlorophyll Fluorescence Response of Persimmon Plants under Salt Stress

[Objectives]To study the photosynthetic response mechanism of persimmon seedlings to salt stress.[Methods]The chlorophyll fluorescence parameters of Diospyros virginiana and Diospyros lotus seedlings under 4%salt stress were studied by pot culture salt control method,including the minimal fluorescence(F_(0)),maximum fluorescence(F_(m)),potential activity of PS II(F_(v)/F_(0)),maximum photochemical efficiency of PS II(F_(v)/F_(m)),electron transport rate(ETR),actual photochemical efficiency of PS II(Y II),and photochemical quenching coefficient(q_(p)).[Results]Under 4%salt stress,the maximum fluorescence(F_(m)),maximum photochemical efficiency of PS II(F v/F m),and photochemical quenching coefficient(q_(p))of two persimmon plants decreased with time.The potential activity of PS II(F_(v)/F_(0)),actual photochemical efficiency of PS II(Y_(II)),and electron transport rate(ETR)decreased under salt stress.[Conclusions]This study indicates that the PS II reaction center in the persimmon leaves was damaged and the electron transport at the acceptor side was damaged under salt stress.It is expected to lay a foundation for the analysis of salt-tolerance mechanism of persimmon plants.

Effect of Exogenous Gibberellin on Wheat Seed Germination under Salt Stress

[Objectives]In this experiment,wheat seeds were treated with different concentrations of gibberellin and different concentrations of salt solution to study the change of germination index of wheat seeds.[Methods]The germination rate,germination potential and germination index of wheat seeds were measured by routine methods,and the effect of exogenous gibberellin on germination of wheat seeds under salt stress was observed.[Results]The germination rate,germination potential and germination index of wheat seeds under salt stress were significantly increased after exogenous treatment of 0.25 and 0.50 g/L gibberellin within the range of salt concentration gradient.However,when the concentration of gibberellin was too high,it would inhibit the germination of seeds.[Conclusions]Appropriate concentration of gibberellin can effectively alleviate the stress caused by salt on wheat seed germination.In this experiment,the best concentration of gibberellin to alleviate salt stress was 0.25 g/L.

Ornithine Pathway in Proline Biosynthesis Activated by Salt Stress in Barley Seedlings

C-14-glutamate and C-14-arginine were spreaded on leaves of six-day old barley (Hordeum vulgare L.) seedlings that were treated with NaCl 200 mmol/L. The result showed that the pathway of arginine-->ornithine-->proline existed in the six-day old barley seedlings and was provoked remarkably by NaCl treatment. After seven days, proline accumulation contributed via the arginine-->ornithine-->proline pathway was 1.0 - 1.5 folds of that via the glutamate-->proline pathway. The activation of arginine-->ornithine-->proline pathway by salt stress in the salt-tolerant cultivar 'Jian 4' was 1.7 - 2.0 folds of that in the salt-sensitive cultivar 'KP 7', which suggested that the activation of arginine-->ornithine-->proline pathway in barley seedlings played an important role in improving salt tolerance of plants.

Cloning and Differential Gene Expression of Two Catalases in Suaeda salsa in Response to Salt Stress

Two different cDNA clones (Sscat1 and Sscat2) encoding catalase, the primary important H2O2-scavenging enzyme, were isolated from a AZap-cDNA library constructed from a 400 mmol/L NaCl-treated library of Suaeda salsa ( L.) Pall aerial tissue. Sscat1 (1.7 kb) contains a full open reading frame of 492 amino acids and Sscat2 (1.1 kb) is a partial clone. BLAST analysis indicates that the two clones share 71.9% identity in nucleotide sequence and 75% identity in deduced amino acid sequence within the last 287 amino acid residues of Sscat1. Southern blotting analysis showed that Sscat1 is multicopy in S. salsa genome, while Sscat2 is a single copy gene. Northern blotting analysis showed a rapid increase in the steady-level of both genes in roots after 48 It salt treatment, but only Sscat1 was induced in salinity treated leaves. Time-course analysis carried out in leaves confirmed that Sscat1 was induced by salt stress, in contrast to Sscat2. These implied that the expression of Sscat1 and Sscat2 genes are differentially regulated in S. salsa. The activity of total catalase is dramatically increased in response to salt stress.

Mechanism of the Effect of Polyamines on the Activity of Tonoplasts of Barley Roots Under Salt Stress

The seeds of barley Hordeum vulgare L. cv. Jian 4) were soaked with 0.1 mmol/L putrescine (Put) and 0.5 mmol/L spermidine (Spd), and then the seedlings were treated with 200 mmol/L NaCl. The growth rate (GR), dry matter accumulation, distribution of ions, the amount of polyamines (PAs) bound to tonoplast proteins as well as lipid composition and the activity of tonoplast vesicles isolated from roots were investigated. The results showed that soaking with Put or Spd could retard salt injury, promote GR and dry matter accumulation, and increase K+/Na+ in the roots. Compared with NaCl_treated plants, phospholipid content in root tonoplast rose by soaking with Put and Spd, while the level of galactose in lipids was decreased. Moreover, the ratio in noncovalently conjugated PA contents of (Spd+PAx (an unknown PA)) to (Put+Dap (diaminopropane)), and the total contents of covalently and noncovalently conjugated PAs were all increased. Statistical analysis indicated that the ratio of (Spd+PAx) to (Put+Dap) was significantly and positively correlated with the activities of membrane associated enzymes H+_ATPase and H+_PPase.

Protective Effects of Glycinebetaine on Brassica chinensis Under Salt Stress

Brassica chinensis L. were foliarly applied with glycinebetaine (GB), as this species is unable to synthesis GB and sensitive to osmotic stress such as salt. The exogenous GB was easily absorbed and transported by the leaf of B. chinensis . Its application (0-20 mmol/L) enhanced the plant tolerance to salt stress. The treatment of 15 mmol/L GB significantly decreased the Na + accumulation in leaf and root under NaCl stress. This difference in accumulating Na + and K + is caused by higher selectivity of root absorption. Furthermore, GB increased H +_ATPase activity of root plasma membrane evidently. This result strongly suggested that in root the decreased Na + accumulation was caused by the GB accumulation that enhanced the extrusion of Na + from the cell in some way through plasma membrane transporter, e.g. Na +/H + antiport driven by H +_ATPase. The GB application was also found to stabilize the plasma membrane, to decrease the loss of chlorophyll, and to stimulate the osmosis induced proline response under salt stress.

Effects of Photoinhibition and Its Recovery on Photosynthetic Functions of Winter Wheat Under Salt Stress

Effects of photoinhibition and its recovery on photosynthetic functions of winter wheat (Triticum aestivum L.) under salt stress were studied. The results showed that several parameters associated with PSⅡ functions, e.g. Fv/Fo?Fv/Fm and qP were not influenced by lower salt concentration (200 mmol/L NaCl) while CO 2 assimilation rate decreased significantly. When exposed to higher salt concentration (400 mmol/L NaCl), PSⅡ functions were significantly inhibited which led to the decrease of carbon assimilation. These results suggest that different concentrations of salt stress affected photosynthesis by different modes. Salt stress made photosynthesis more sensitive to strong light and led to more serious photoinhibition. Under lower concentration of salt stress, the Q B-non-reductive PSⅡ reaction centers formed at the beginning of photoinhibition could be effectively used to compose active PSⅡ reaction center (RC) and repair the reversible inactivated PSⅡ RC. Under higher concentration of salt stress, PSⅡ reaction centers were seriously damaged during photoinhibition, the Q B-non-reductive PSⅡ RC could only be partly effective at the early time of photoinhibition, thus led to the accumulation of Q B-non-reductive PSⅡ RC in the course of restoration under dim light.

Changes in Physiological Properties and Respiratory Pathway of the New Lines of Wheat Introduced Exogenous DNA Under Salt Stress

New lines of wheat ( Triticum aestivum L.) was obtained by introducing the DNA of sorghum (Sorghum vulgare Pers.) into wheat cultivar 'Longchun 13'. The changes of respiratory pathway, contents of protein, Na+ and K+ in the leaves and roots of the new lines of wheat under salt stress were determined and compared with the control cultivar, 'Longchun 13'. The decrease of the content of K+ was observed with the increase of NaCl concentrations, but the decrease was more in the control than that in the new lines, and more in roots than in leaves. Content of proline and Na+ in both two wheats lines increased greatly, but the former increased more significantly in the new lines and the latter more significantly in control both in leaves and roots. The operation of the cyanide-resistant pathway of respiration was enhanced at different degrees after salt stress and it increased much more in roots and leaves of the control plant than that in the new lines, but the cytochrome pathway of electron transport was still the main one consistently. The possible significance of these changes was discussed.

Effect of Exogenous GSH on Growth of Barley Seedlings under Salt Stress

The growth of barley (Hordeum vulgare L.) seedling is inhibited by 300 mmol/L NaCl. When 20 mg/L GSH is present in the hydroponic culture solution with NaCl, root length, root and shoot dry weigh, chlorophyll as well as K+ contents are increased, Na+ content is decreased. At the same time the activities of H+-ATPase and H+-PPase associated with tonoplast vesicles isolated from leaves are stimulated, and electrolytic leakage are diminished by exogenous GSH.

Effects of Polyamines on Plants under Salt Stress

Salt stress is one of the worldwide abiotic stresses resulting in growth re- duction, chlorosis, wilting, and plant death. These exhibitions might result from men- tal toxicity and osmotic stress induced by salt. The two aspects of stress would af- fect vital metabolic pathways, reactive oxygen species scavenging system, lipid per- oxidation and photosynthetic apparatus. Thus, exploring ways to improve crop pro- ductivity or alleviate harmful effects under salt is one of the major areas of concern. Polyamines are aliphatic nitrogen organic cations which are implicated in a wide range of plant physiological processes such as morphogenesis, flower differentiation and initiation, they also play a role in biotic or abiotic stress responses. At the physiological level, polyamines modify the activities of many enzymes included in salt stress response and can bond to photosynthetic apparatus, thus changing the photosynthetic efficiency. At molecular level, polyamines can modify expressions of the polyamine-related genes directly or indirectly. Significant researches had been done to understand the effects of polyamines on plant salt resistance, but several questions still need to be answered. The present review is focused specifically on the effects of polyamines on physiological and molecular changes in plants under salt stress.