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Salt stress is one of the significant environmental stressors that severely affects plant growth and development. Plant responses to salt stress involve a series of biological mechanisms, including osmoregulation, redox and ionic homeostasis regulation, as well as hormone or light signaling-mediated growth adjustment, which are regulated by different functional components. Unraveling these adaptive mechanisms and identifying the critical genes involved in salt response and adaption are crucial for developing salt-tolerant cultivars. This review summarizes the current research progress in the regulatory networks for plant salt tolerance, highlighting the mechanisms of salt stress perception, signaling, and tolerance response. Finally, we also discuss the possible contribution of microbiota and nanobiotechnology to plant salt tolerance.

Soil salinization is a widespread hindrance that endangers agricultural production and ecological security. High salt concentrations in saline soils are primarily caused by osmotic stress, ionic toxicity and oxidative stress, which have a negative impact on plant growth and development. In order to withstand salt stress, plants have developed a series of complicated physiological and molecular mechanisms, encompassing adaptive changes in the structure and function of various plant organs, as well as the intricate signal transduction networks enabling plants to survive in high-salinity environments. This review summarizes the recent advances in salt perception under different tissues, physiological responses and signaling regulations of plant tolerance to salt stress. We also examine the current knowledge of strategies for breeding salt-tolerant plants, including the applications of omics technologies and transgenic approaches, aiming to provide the basis for the cultivation of salt-tolerant crops through molecular breeding. Finally, future research on the application of wild germplasm resources and muti-omics technologies to discover new tolerant genes as well as investigation of crosstalk among plant hormone signaling pathways to uncover plant salt tolerance mechanisms are also discussed in this review.

In animal cells, phospholipase C (PLC) isoforms predominantly hydrolyze phosphatidylinositol-4,5-biphosphates [PtdIns(4,5)P2] into the second messengers diacylglycerol (DAG) and inositol 1,4,5-trisphosphate [Ins(1,4,5)P3] to regulate diverse biological processes. By contrast, the molecular mechanisms and physiological significance of PLC signaling in plants still awaits full elucidation. Here, we identified a rice (Oryza sativa cv) PI-PLC, OsPLC1, which preferred to hydrolyze phosphatidylinositol-4-phosphate (PtdIns4P) and elicited stress-induced Ca2+ signals regulating salt tolerance. Analysis by ion chromatography revealed that the concentration of PtdIns4P was c. 28 times of that of PtdIns(4,5)P2 in shoots. OsPLC1 not only converted PtdIns(4,5)P2 but also – and even more efficiently – converted PtdIns4P into DAG and Ins(1,4,5)P3 in vitro and in vivo. Salt stress induced the recruitment of OsPLC1 from cytoplasm to plasma membrane, where it hydrolyzed PtdIns4P. The stress-induced Ca2+ signaling was dependent on OsPLC1, and the PLC-mediated Ca2+ signaling was essential for controlling Na+ accumulation in leaf blades, thus establishing whole plant salt tolerance. Our work identifies a conversion pathway and physiological function for PtdIns4P pools in rice and reveals the connection between phosphoinositides and Ca2+ signals mediated by PLC during salt stress responses.

Salt is a major environmental stress factor that can affect rice growth and yields. Recent studies suggested that members of the AP2/ERF domain-containing RAV (related to ABI3/VP1) TF family are involved in abiotic stress adaptation. However, the transcriptional response of rice RAV genes (OsRAVs) to salt has not yet been fully characterized. In this study, the expression patterns of all five OsRAVs were examined under salt stress. Only one gene, OsRAV2, was stably induced by high-salinity treatment. Further expression profile analyses indicated that OsRAV2 is transcriptionally regulated by salt, but not KCl, osmotic stress, cold or ABA (abscisic acid) treatment. To elucidate the regulatory mechanism of the stress response at the transcriptional level, we isolated and characterized the promoter region of OsRAV2 (P OₛRAV₂). Transgenic analysis indicated that P OₛRAV₂ is induced by salt stress but not osmotic stress or ABA treatment. Serial 5′ deletions and site-specific mutations in P OₛRAV₂ revealed that a GT-1 element located at position −664 relative to the putative translation start site is essential for the salt induction of P OₛRAV₂ . The regulatory function of the GT-1 element in the salt induction of OsRAV2 was verified in situ in plants with targeted mutations generated using the CRISPR/Cas9 (clustered regularly interspaced short palindromic repeats/CRISPR-associated protein 9) system. Taken together, our results indicate that the GT-1 element directly controls the salt response of OsRAV2. This study provides a better understanding of the putative functions of OsRAVs and the molecular regulatory mechanisms of plant genes under salt stress.

Key message The study on melatonin biosynthesis mutant snat1snat2 revealed that endogenous melatonin plays an important role in salt responsiveness by mediating auxin signaling. Melatonin is a pleiotropic signaling molecule, which, besides being involved in multiple growth and developmental processes, also mediates environmental stress responses. However, whether and how endogenous melatonin is involved in salt response has not been determined. In this study, we elucidated the involvement of endogenous melatonin in salt response by investigatiing the impact of salt stress on a double mutant of Arabidopsis ( snat1snat2 ) defective in melatonin biosynthesis genes SNAT1 and SNAT2 . This mutant was found to exhibit salt sensitivity, manifested by unhealthy growth, ion imbalance and ROS accumulation under salt stress. Transcriptomic profiles of snat1snat2 revealed that the expression of a large number of salt-responsive genes was affected by SNAT defect, and these genes were closely related to the synthesis of auxin and several signaling pathways. In addition, the salt-sensitive growth phenotype of snat1snat2 was alleviated by the application of exogenous auxin. Our results show that endogenous melatonin may be essential for plant salt tolerance, a function that could be correlated with diverse activity in mediating auxin signaling.

Clarifying the morphological, physiological and molecular mechanisms of rice seedlings in response to salt stress is essential for improving salt tolerance and rice yield. Here, we systematically analyzed the responses of Nipponbare seedlings to salt stress using a two-stage approach. First, NaCl concentration treatments (0–170 mM) revealed a concentration-dependent inhibition of seedling growth. Then, NaCl treatments for 0, 6, and 12 h at the critical concentration (90 mM) were conducted to characterize short-term physiological responses. Transcriptome analysis showed temporally specific transcriptional patterns, with 20 salt stress-related co-expression modules identified. Notably, the MEblue module correlated strongly with superoxide dismutase (SOD) ( r  = 0.93) and peroxidase (POD) ( r  = 0.9) activities, while the MEpink module was strongly correlated with H 2 O 2 content ( r  = 0.92). Core hub genes ( OsSAMS1 , ADK2 , et al.) were screened from these modules. This study established the critical salt-tolerance concentration of Nipponbare seedlings, characterized their dynamic physiological responses to short-term salt stress, identified the primary antioxidant pathways, and screened potential regulatory genes, thereby providing gene targets and a theoretical basis for understanding salt tolerance mechanisms in rice seedlings.

Key messageConstruction of ML-hGRN for the salt pathway in Populus davidiana × P. bolleana. Construction of ML-hGRN for the lignocellulosic pathway in Populus davidiana × P. bolleana under salt stress.Many woody plants, including Populus davidiana × P. bolleana, have made great contributions to human production and life. High salt is one of the main environmental factors that restricts the growth of poplar. This study found that high salt could induce strong biochemical changes in poplar. To detect the effect of salt treatment on gene expression, 18 libraries were sequenced on the Illumina sequencing platform. The results identified a large number of early differentially expressed genes (DEGs) and a small number of late DEGs, which indicated that most of the salt response genes of poplar were early response genes. In addition, 197 TFs, including NAC, ERF, and other TFs related to salt stress, were differentially expressed during salt treatment, which indicated that these TFs may play an important role in the salt stress response of poplar. Based on the RNA-seq analysis results, multilayered hierarchical gene regulatory networks (ML-hGRNs) of salt stress- and lignocellulosic synthesis-related DEGs were constructed using the GGM algorithm. The lignocellulosic synthesis regulatory network under salt stress revealed that lignocellulosic synthesis might play an important role in the process of salt stress resistance. Furthermore, the NAC family transcription factor PdbNAC83, which was found in the upper layer in both pathways, was selected to verify the accuracy of the ML-hGRNs. DAP-seq showed that the binding site of PdbNAC83 included a “TT(G/A)C(G/T)T” motif, and ChIP-PCR further verified that PdbNAC83 can regulate the promoters of at least six predicted downstream genes (PdbNLP2-2, PdbZFP6, PdbMYB73, PdbC2H2-like, PdbMYB93-1, PdbbHLH094) by binding to the “TT(G/A)C(G/T)T” motif, which indicates that the predicted regulatory network diagram obtained in this study is relatively accurate. In conclusion, a species-specific salt response pathway might exist in poplar, and this finding lays a foundation for further study of the regulatory mechanism of the salt stress response and provides new clues for the use of genetic engineering methods to create high-quality and highly resistant forest germplasms.

Euphrates poplar (Populus euphratica) is known as a system model to study the genomic mechanisms underlying the salt resistance of woody species. To characterize how dynamic gene regulatory networks (GRNs) drive the defense response of this species to salt stress, we performed mRNA sequencing of P. euphratica roots under short-term (ST) and long-term (LT) salt stress treatments across multiple time points. Comparisons of these transcriptomes revealed the diverged gene expression patterns between the ST and LT treated samples. Based on the informative, dynamic, omnidirectional, and personalized networks model (idopNetwork), inter- and intra-module networks were constructed across different time points for both the ST and LT groups. Through the analysis of the inter-module network, we identified module 4 as the hub, containing the largest number of genes. Further analysis of the gene network within module 4 revealed that gene XM_011048240.1 had the most prominent interactions with other genes. Under short-term salt stress, gene interactions within the network were predominantly promoted, whereas under long-term stress, these interactions shifted towards inhibition. As for the gene ontology (GO) annotation of differentially expressed genes, the results suggest that P. euphratica may employ distinct response mechanisms during the early and late stages of salt stress. Taking together, these results offer valuable insights into the regulatory mechanism involved in P. euphratica’s stress response, advancing our understanding of complex biological processes.

Cytochrome P450 (CYP450) proteins are a large group of monooxygenase that play important roles in the biosynthesis of secondary metabolites and degradation of xenobiotics. However, the responses of CYP450 family to abiotic stresses have not been characterized in Brassica napus ( B. napus ). In this study, we identified a total of 384 CYP450 genes in Darmor- bzh , the rapeseed culture whose genome was wildly used as a reference for gene clone. The structure and localization analyses showed that BnaCYP450 genes have integrated heme-binding motif, contain 1–10 exons, unevenly distributed across all the 19 chromosomes, and mainly localized on chloroplast. Cis-regulation element analysis suggested that BnaCYP450 genes were transcriptionally regulated by hormone and multiple stress response signals. Transcript expression analyses identified 108, 85, 96, and 86 BnaCYP450s differentially expressed genes (DEGs) in response to salt stress, potassium deficiency, nitrogen stress, and cadmium toxicity, respectively. Gene ontology (GO) enrichment analysis indicated that these BnaCYP450 DEGs mainly enriched in molecular function of ion binding and oxidoreductase activity and the biological process of secondary product metabolism. Kyoto Encyclopedia of Genes and Genomes (KEGG) enrichment analysis showed that they mainly involved in the pathway of isoflavonoid biosynthesis. Differential expression of BnaCYP450s to multiple abiotic stresses revealed the functional diversity of BnaCYP450 family in B. napus . This study gave a basic understanding of CYP450 genes in B. napus and provides multiple core BnaCYP450 genetic resources for improving plant resistance to multiple abiotic stresses.

Jasmonates (JAs) together with jasmonic acid and its offshoots are lipid-derived endogenous hormones that play key roles in both developmental processes and different defense responses in plants. JAs have been studied intensively in the past decades for their substantial roles in plant defense comebacks against diverse environmental stresses among model plants. However, the role of this phytohormone has been poorly investigated in the monocotyledonous species against abiotic stresses. In this study, a JA biosynthesis mutant opr7opr8 was used for the investigation of JA roles in the salt stress responses of maize seedlings, whose roots were exposed to 0 to 300 mM NaCl. Foliar stomatal observation showed that opr7opr8 had a larger stomatal aperture than wild type (WT) (B73) under salinity stress, indicating that JA positively regulates guard cell movement under salt stress. The results regarding chlorophyll content and leaf senescence showed that opr7opr8 exhibited delayed leaf senescence under salt stress as compared to WT, indicating that JA plays a role in salt-inducing cell death and subsequent leaf senescence. Moreover, the morphological parameters, including the length of the shoots and roots, and the fresh and dry weights of the shoots and roots, showed that after 7 days of salt treatment, opr7opr8 had heavier and longer shoots than WT but slighter and shorter roots than WT. In addition, ion analysis showed that opr7opr8 accumulated less sodium but more potassium in the leaves than WT but more sodium and less potassium in the roots than WT, suggesting that JA deficiency causes higher salt stress to the roots but less stress to the leaves of the seedlings. Reactive oxygen species (ROS) analysis showed that opr7opr8 produced less H2O2 than WT in the leaves but more H2O2 in the roots under salt treatment, and correspondingly, ROS-scavenging enzymes superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX) showed a similar variation, i.e., opr7opr8 has lower enzymatic activities in the shoots but higher activities in the roots than WT under salt treatment. For osmotic adjustment, opr7opr8 produced less proline in the shoots at 100 and 300 mM NaCl treatments but more in the roots than the WT roots under all salt treatments. In addition, the gene expression for abscisic acid (ABA) biosynthesis under salt stress was investigated. Results showed that the expression levels of four key enzymes of ABA biosynthesis, ZEP1, NCED5, AO1, and VP10, were significantly downregulated in the shoots as compared to WT under salt treatment. Putting all the data together, we concluded that JA-deficiency in maize seedlings reduced the salt-stress responses in the shoots but exaggerated the responses in the roots. In addition, endogenous JA acted as a positive regulator for the transportation of sodium ions from the roots to the shoots because the mutant opr7opr8 had a higher level of sodium in the roots but a significantly lower level in the shoots than WT. Furthermore, JA may act as a positive regulator for ABA biosynthesis in the leaves under salt stress.