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      Identifying resurrection genes through the differentially expressed genes between Selaginella tamariscina (Beauv.) spring and Selaginella moellendorffii Hieron under drought stress

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          Abstract

          Selaginella tamariscina (Beauv.) spring, a primitive vascular resurrection plant, can survive extreme drought and recover when water becomes available. To identify drought-inducible genes and to clarify the molecular mechanism of drought tolerance, a comparative transcriptional pattern analysis was conducted between S. tamariscina and Selaginella moellendorffii Hieron (drought sensitive). 133 drought related genes were identified, including 72 functional genes and 61 regulatory genes. And several drought responsive reactions, such as antioxidant activity, osmotic balance, cuticle defense and signal transduction were highlighted in S. tamariscina under drought. Notably, besides peroxidase, catalase and L-ascorbate oxidase genes, DEGs associated with phenylalanine metabolism and polyamine catabolism could be alternative ways to enhance antioxidant ability in S. tamariscina. DEGs related to soluble carbohydrate metabolism, late embryogenesis abundant protein (LEA) and aquaporin protein (AQP) confirmed that osmotic adjustment could resist drought during desiccation. DEGs involved in xyloglucan metabolic process, pectin metabolic process and cutin biosynthesis may also contribute to drought tolerance of S. tamariscina by cuticle defense. Drought-responsive genes encoding protein kinases, calcium sensors, transcription factors (TFs) and plant hormones also help to drought resistance of S. tamariscina. The preliminary validation experiments were performed and the results were consistent with our hypothetical integrated regulatory network. The results of this study provide candidate resurrection genes and an integrated regulatory network for further studies on the molecular mechanisms of stress tolerance in S. tamariscina.

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          Response of Phenylpropanoid Pathway and the Role of Polyphenols in Plants under Abiotic Stress

          Phenolic compounds are an important class of plant secondary metabolites which play crucial physiological roles throughout the plant life cycle. Phenolics are produced under optimal and suboptimal conditions in plants and play key roles in developmental processes like cell division, hormonal regulation, photosynthetic activity, nutrient mineralization, and reproduction. Plants exhibit increased synthesis of polyphenols such as phenolic acids and flavonoids under abiotic stress conditions, which help the plant to cope with environmental constraints. Phenylpropanoid biosynthetic pathway is activated under abiotic stress conditions (drought, heavy metal, salinity, high/low temperature, and ultraviolet radiations) resulting in accumulation of various phenolic compounds which, among other roles, have the potential to scavenge harmful reactive oxygen species. Deepening the research focuses on the phenolic responses to abiotic stress is of great interest for the scientific community. In the present article, we discuss the biochemical and molecular mechanisms related to the activation of phenylpropanoid metabolism and we describe phenolic-mediated stress tolerance in plants. An attempt has been made to provide updated and brand-new information about the response of phenolics under a challenging environment.
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            Functional analysis of an Arabidopsis transcription factor, DREB2A, involved in drought-responsive gene expression.

            Transcription factors DREB1A/CBF3 and DREB2A specifically interact with cis-acting dehydration-responsive element/C-repeat (DRE/CRT) involved in cold and drought stress-responsive gene expression in Arabidopsis thaliana. Intact DREB2A expression does not activate downstream genes under normal growth conditions, suggesting that DREB2A requires posttranslational modification for activation, but the activation mechanism has not been clarified. DREB2A domain analysis using Arabidopsis protoplasts identified a transcriptional activation domain between residues 254 and 335, and deletion of a region between residues 136 and 165 transforms DREB2A to a constitutive active form. Overexpression of constitutive active DREB2A resulted in significant drought stress tolerance but only slight freezing tolerance in transgenic Arabidopsis plants. Microarray and RNA gel blot analyses revealed that DREB2A regulates expression of many water stress-inducible genes. However, some genes downstream of DREB2A are not downstream of DREB1A, which also recognizes DRE/CRT but functions in cold stress-responsive gene expression. Synthetic green fluorescent protein gave a strong signal in the nucleus under unstressed control conditions when fused to constitutive active DREB2A but only a weak signal when fused to full-length DREB2A. The region between DREB2A residues 136 and 165 plays a role in the stability of this protein in the nucleus, which is important for protein activation.
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              A Raf-like MAPKKK gene DSM1 mediates drought resistance through reactive oxygen species scavenging in rice.

              Mitogen-activated protein kinase (MAPK) cascades have been identified in various signaling pathways involved in plant development and stress responses. We identified a drought-hypersensitive mutant (drought-hypersensitive mutant1 [dsm1]) of a putative MAPK kinase kinase (MAPKKK) gene in rice (Oryza sativa). Two allelic dsm1 mutants were more sensitive than wild-type plants to drought stress at both seedling and panicle development stages. The dsm1 mutants lost water more rapidly than wild-type plants under drought stress, which was in agreement with the increased drought-sensitivity phenotype of the mutant plants. DSM1-RNA interference lines were also hypersensitive to drought stress. The predicted DSM1 protein belongs to a B3 subgroup of plant Raf-like MAPKKKs and was localized in the nucleus. By real-time PCR analysis, the DSM1 gene was induced by salt, drought, and abscisic acid, but not by cold. Microarray analysis revealed that two peroxidase (POX) genes, POX22.3 and POX8.1, were sharply down-regulated compared to wild type, suggesting that DSM1 may be involved in reactive oxygen species (ROS) signaling. Peroxidase activity, electrolyte leakage, chlorophyll content, and 3,3'-diaminobenzidine staining revealed that the dsm1 mutant was more sensitive to oxidative stress due to an increase in ROS damage caused by the reduced POX activity. Overexpression of DSM1 in rice increased the tolerance to dehydration stress at the seedling stage. Together, these results suggest that DSM1 might be a novel MAPKKK functioning as an early signaling component in regulating responses to drought stress by regulating scavenging of ROS in rice.
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                Author and article information

                Contributors
                Role: Formal analysisRole: MethodologyRole: Project administrationRole: ValidationRole: Writing – original draftRole: Writing – review & editing
                Role: Formal analysisRole: Writing – original draftRole: Writing – review & editing
                Role: Project administrationRole: Writing – review & editing
                Role: Project administrationRole: Writing – review & editing
                Role: Project administrationRole: Writing – original draftRole: Writing – review & editing
                Role: Formal analysisRole: Writing – review & editing
                Role: SupervisionRole: Writing – review & editing
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                13 November 2019
                2019
                : 14
                : 11
                : e0224765
                Affiliations
                [1 ] School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing, China
                [2 ] Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing, China
                Chinese Academy of Medical Sciences and Peking Union Medical College, CHINA
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Author information
                http://orcid.org/0000-0001-7109-7848
                Article
                PONE-D-19-17762
                10.1371/journal.pone.0224765
                6853609
                31721818
                95956875-c688-4ceb-a58a-ca7c812b8f8c
                © 2019 Gu 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.

                History
                : 23 June 2019
                : 21 October 2019
                Page count
                Figures: 6, Tables: 1, Pages: 20
                Funding
                Funded by: Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization
                Award ID: Grant No. ZDXM-HT-1-5 and ZDXM-3-24
                Award Recipient :
                Funded by: Priority Academic Program Development of Jiangsu Higher Education Institutions (CN)
                Award ID: ysxk-2014
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, 2017 Chinese Medicine Public Health Service Subvention Special "National traditional Chinese Medicine Resource Survey Project";
                Award ID: Caishe (2017) 66
                Award Recipient :
                This research was supported by Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization (Grant No. ZDXM-HT-1-5 and ZDXM-3-24), the Priority Academic Program Development of Jiangsu Higher Education Institutions (ysxk-2014) and 2017 Chinese Medicine Public Health Service Subvention Special "National traditional Chinese Medicine Resource Survey Project" (Caishe (2017) 66). Jin-Ao Duan was the host of the above-mentioned four projects. This research was also supported by the National Natural Science Foundation of China(Grant No. 81573520) and Jianguo Chao was the host of this project. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Genetics
                Gene Expression
                Biology and Life Sciences
                Plant Science
                Plant Physiology
                Plant Defenses
                Plant Resistance to Abiotic Stress
                Biology and Life Sciences
                Plant Science
                Plant Pathology
                Plant Resistance to Abiotic Stress
                Biology and Life Sciences
                Ecology
                Plant Ecology
                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Ecology and Environmental Sciences
                Ecology
                Plant Ecology
                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Biology and Life Sciences
                Plant Science
                Plant Ecology
                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Medicine and Health Sciences
                Diagnostic Medicine
                Signs and Symptoms
                Dehydration (Medicine)
                Medicine and Health Sciences
                Pathology and Laboratory Medicine
                Signs and Symptoms
                Dehydration (Medicine)
                Ecology and Environmental Sciences
                Natural Resources
                Water Resources
                Biology and Life Sciences
                Plant Science
                Plant Physiology
                Plant Defenses
                Plant Resistance to Abiotic Stress
                Drought Adaptation
                Biology and Life Sciences
                Plant Science
                Plant Pathology
                Plant Resistance to Abiotic Stress
                Drought Adaptation
                Biology and Life Sciences
                Ecology
                Plant Ecology
                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Drought Adaptation
                Ecology and Environmental Sciences
                Ecology
                Plant Ecology
                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Drought Adaptation
                Biology and Life Sciences
                Plant Science
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                Plant-Environment Interactions
                Plant Resistance to Abiotic Stress
                Drought Adaptation
                Physical Sciences
                Chemistry
                Chemical Compounds
                Organic Compounds
                Carbohydrates
                Disaccharides
                Trehalose
                Physical Sciences
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                Organic Compounds
                Carbohydrates
                Disaccharides
                Trehalose
                Biology and Life Sciences
                Biochemistry
                Enzymology
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                Biology and Life Sciences
                Biochemistry
                Proteins
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                Biology and Life Sciences
                Biochemistry
                Antioxidants
                Custom metadata
                The raw data of S. tamariscina and S. moellendorfii were submitted to the Short Read Archive database of National Center for Biotechnology Information (NCBI) under the project accession number SRR8149073 and SRR8149074, and in the fastq file format.

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