Glyma11g13220, a homolog of the vernalization pathway gene VERNALIZATION 1 from soybean [Glycine max (L.) Merr. The contrasting ecologies of these three species thus predict major consequences on their strategies to face up to the challenges imposed by water limitations. PLoS One 11:e157026. However, eukaryotic organisms are complex and a certain signal transduction pathway for some intermediates can at the same time be involved in other physiological and biochemical reactions or other metabolic pathways in the cell. In contrast to rice and Arabidopsis, many WRKY genes in other plants have been identified specifically in relation to abiotic stress. The primers used for qRT-PCR are listed in Table S1. in transgenic rice plants (Tables 1, 2). To investigate the tolerance to osmotic stress of GmWRKY16 transgenic lines after molecular identification (Supplementary Figure S1), different treatments of mannitol and NaCl were performed on 1/2 MS medium to determine the germination rates and root elongation of Arabidopsis (Figures 4, 5). Jonak C, Hirt H. Glycogen synthase kinase 3/SHAGGY-like kinases in plants: an emerging family with novel functions. doi: 10.1007/s00425-011-1564-z, Liu, L., Hu, X., Song, J., Zong, X., Li, D., and Li, D. (2009). Among the WRKY genes in soybean, the overexpression of GmWRKY21 enhanced the tolerance to cold stress in Arabidopsis, whereas GmWRKY54 conferred salt and drought tolerance. Barplots with one asterisk or more are significantly different (Tukeys HSD, P < 0.1; ***P < 0.001; ns, not significant). The ABI1 and ABI2 protein phosphatases 2C act in a negative feedback regulatory loop of the abscisic acid signalling pathway. doi: 10.1093/molbev/msw054, Li, J. The list and expression data of up-regulated genes in response to drought and salinity stress are available as a supplementary material (Table S1). Zhang T, Tan D, Zhang L, Zhang X, Han Z. Phylogenetic analysis and drought-responsive expression profiles of the WRKY transcription factor family in maize. The genome-wide expression profiles of Hsfs genes in B. napus revealed that many members of BnaHsf family respond to drought stress. Department of Biology, Appalachian State University, Boone, North Carolina, United States of America, Affiliation: (2014). doi: 10.1371/journal.pone.0157026, Clough, S. J., and Bent, A. F. (1998). Stomatal density and 13C measured in Arabidopsis halleri and A. lyrata grown under well-watered conditions. Front. These results indicated that GmWRKY16 protein had transcriptional activity in yeast cell. Plant Physiol. 25, 239250. These results suggest that overexpression of GmWRKY16 enhances the Arabidopsis tolerance to ABA stress. (2015). doi: 10.1046/j.1365-313x.1998.00343.x, Coue, I., Sulmon, C., Gouesbet, G., and El Amrani, A. College of Life Science, Zhejiang University, Hangzhou, Zhejiang, PR China, Affiliation: Table S12: differentially expressed genes identified for each of Arabidopsis halleri and A. lyrata between 20 and 60 % soil moisture and between recovery and 60 % soil moisture. Annu Rev Plant Physiol Plant Mol Biol 2000; 51(1): 463-99. ", "Open access journals are a novel concept in the medical literature. Seeds of Arabidopsis ecotype Columbia (Col-0) were used in this study. 1). Plantarum 126, 519527. Expressional analysis and role of calcium regulated kinases in abiotic stress signaling. doi: 10.1111/j.1467-7652.2011.00634.x, Rushton, P. J., Somssich, I. E., Ringler, P., and Shen, Q. J. (C,D) Seedlings under the treatment of salt. The 3-week-old Arabidopsis plants were then irrigated with a solution of 200 mM NaCl for 15 days (Niu et al., 2012; Ying et al., 2012; Babitha et al., 2013). Yet, the absence of a significant interaction term between initial rosette area and species (M4: F2, 170 = 1.89, P-value = 0.15) indicated that interspecific differences in plant size did not explain interspecific differences in the rate of soil water consumption. Overexpression of VaWRKY14 increases drought tolerance in Arabidopsis by modulating the expression of stress-related genes. The list and expression data on these drought and salinity stress-down-regulated gene accessions are available as a supplementary material (Table S2). Plant. The result of CARE analysis demonstrated that the light responsive elements were mainly controlled by the responsive genes. There are different products of genes with kinase activity, which typically makes them useful for abiotic stress tolerance. EMBO Rep. 14, 11361142. Collectively, our data show that SF priming mitigates multiple cellular processes that otherwise impair plant growth under drought stress, thereby providing a knowledge basis for future research on crops. 61, 25192533. Acta 1819:85. doi: 10.1016/j.bbagrm.2012.01.005. As part of our efforts to improve tomato tolerance to abiotic stress, we have undertaken this study to introduce two candidate genes encoding: a sodium antiporter and a vacuolar pyrophosphatase, previously shown to enhance drought and salt tolerance in transgenic Arabidopsis plants. These two accessions are representative of the phenotypic diversity observed in the dry-down experiment. Under stressful conditions, transcriptome changes are the earliest responses in plants. (2012). The full-flowering Arabidopsis plants were used for genetic transformation by the floral dip method (Clough and Bent, 1998). One to two weeks later, survival and symptoms of damage were scored. ", "Open access journals are freely available online throughout the world, for you to read, download, copy, distribute, and use. According to the results of GO analysis, 21 genes including HSFC1, AT3G24518, CID12, CID11, GATA10, GATA11, APC1, APC3, RHC2A, PDCB4, APRR2, GRP2, GRP2B, psaC, AT5G10770, NBR1, AT1G13190, AT5G55670, TIM23-1, TIM23-2 and TIM23-3 had significant role in binding activity as TF elements. Thomashow MF. Future work will have to investigate the impact of modifications in stomatal patterning on interspecific differences in tolerance and avoidance in the face of limiting SWC. In order to gain insight into the molecular changes underpinning these differences, we performed a third dry-down experiment to collect leaf material in one representative accession of each of the sister species A. halleri and A. lyrata, and examined the reaction to stress and recovery at the transcriptome level. The random expectation of percentage overlap is indicated in the top row. Error bars represent SD. The results of t-test for the microarray data demonstrated 2558 and 3691 gene accessions were differentially expressed in response to drought stress in root and shoot tissue, respectively. The young leaves of 4-week-old tobacco plants were contaminated by the recombinant Agrobacterium tumefaciens using the method according to Kokkirala et al. DREBs (dehydration responsive element binding) are important plant Transcription Factors (TFs) regulating the expression of many stress-inducible genes mostly in an ABA-independent manner and play a critical role in improving the abiotic stress tolerance of plants by interacting with a DRE/CRT cis-element present in the promoter region of various abiotic stress-responsive genes [56]. We are very grateful to Prof. Yaoguang Liu (South China Agricultural University, China) for kindly providing us with the pCAMBIA1302 vector. The relative importance of strategies to cope with drought stress is expected to be intimately linked to the life history and ecology of species. RNA extraction was performed using the PureLink RNA Ambion Mini Kit (Thermofisher, Darmstadt, Germany). A moso bamboo WRKY gene PeWRKY83 confers salinity tolerance in transgenic Arabidopsis plants. The inner reference gene -tubulin was used to normalize the data. Overexpression of SpWRKY1 promotes resistance to Phytophthora nicotianae and tolerance to salt and drought stress in transgenic tobacco. The partnership allows the researchers from the university to publish their research under an Open Access license with specified fee discounts. The results indicated that 0.25% (27 out of 10645) of DEGs from both tissues showed possible crosstalk in response to drought and salinity stresses (Table 2). *Correspondence: Hai Nian, hnian@scau.edu.cn, These authors have contributed equally to this work, https://www.frontiersin.org/articles/10.3389/fpls.2018.01979/full#supplementary-material, Creative Commons Attribution License (CC BY). An individual WRKY gene can enhance plant tolerance to abiotic stress (Wang et al., 2013; Bakshi and Oelmller, 2014; Dai et al., 2016; Kiranmai et al., 2018). The articles are high standard and cover a wide area. Wilting manifested differently in the three species. doi: 10.1104/pp.112.202143, Seo, Y., Park, J., Cho, Y., Jung, C., Seo, H., Park, S., et al. p-values were adjusted in false discovery rate as less than <0.05. All the GmWRKY16 transgenic seedlings were screened by an herbicide to obtain the positive plants. Xiong L, Schumaker KS, Zhu JK. Its specific ability to accumulate heavy metals enhances its defences against herbivores but sets strong constitutive demands on detoxifying systems which are important for re-establishing homeostasis after stress (Mittler, 2002; Becher et al., 2004; Krmer and Clemens, 2006; Stolpe et al., 2016). Bookshelf By comparison, a recent and exhaustive analysis of stomatal density in A. thaliana reported that stomatal density varies from 87 to 204 stomata mm2 and it is negatively correlated with stomatal length (Dittberner et al., 2018). BMC Plant Biol. For each species, we compared transcript abundance at three time points during the dry-down experiment, i.e. Curr Opin Plant Biol 2002; 5(5): 408-14. Sulfur dioxide (SO 2 ) is a common air pollutant that has multiple effects on plants. Front. In recent years, numerous data linking cellular responses and regulation of gene expression have been collected, especially on the model plant Arabidopsis thaliana. SpringerPlus 5:920. doi: 10.1186/s40064-016-2647-x, Zeng, Q., Yang, C., Ma, Q., Li, X., Dong, W., and Nian, H. (2012). Wan J, Dunning FM, Bent AF. Greenham K, McClung CR. Comparative Transcriptome Analysis of Two Ascophyllum nodosum Extract Biostimulants: Same Seaweed but Different. In this paper we have shown that the expression of MIR398a and MIR398c/MIR398b first increased as PEG8000 concentration went up from 0 to 0.05 g ml1/0.2 g ml1 and then decreased (Figure 2B). Chrysanthemum WRKY gene DgWRKY5 enhances tolerance to salt stress in transgenic chrysanthemum. All the drought-treated plants were then rehydrated after they were dried for 5 days. doi: 10.1016/j.agwat.2018.01.028. The maize WRKY transcription factor ZmWRKY40 confers drought resistance in transgenic Arabidopsis. Subcellular localization of proteins of Oryza sativa L. in the model tobacco and tomato plants. doi: 10.1093/mp/sss080, Jung, C., Seo, J. S., Han, S. W., Koo, Y. J., Kim, C. H., Song, S. I., et al. The positive clones in Escherichia coli were used to obtain the full cDNA sequence of GmWRKY16 identified by PCR, enzyme digestion and sequencing [Sangon Biotech (Shanghai) Co., Ltd., China] (L et al., 2015; Ma et al., 2018). The HVA1 transgenic rice plants had showed significantly increased tolerance to water deficit and salinity [40]. When lcd mutants (with lower H2S production rate than WT) were treated with PEG8000, they showed lower levels of miRNA expression changes than WT. The taken samples of Arabidopsis were frozen immediately in liquid nitrogen and stored at -80C (Chen et al., 2015). 9:346. doi: 10.3389/fpls.2018.00346, Kokkirala, V. R., Yonggang, P., Abbagani, S., Zhu, Z., and Umate, P. (2010). Yu LH, Wu SJ, Peng YS, Liu RN, Chen X, Zhao P, Xu P, Zhu JB, Jiao GL, Pei Y, Xiang CB., Plant Biotechnol J 14(1), 2016 PMID: 25879154. Molecular characteriza-tion of novel TaNAC genes in wheat and overexpres-sion of TaNAC2a confers drought tolerance in tobac-co. Physiol Plant. Among genes, the role of AT3G24518 was remarkable because it codes a non-coding RNA which suggests some possible epigenetic regulation mechanisms like histone modification and chromatin remodeling. CSD1and CSD2 (targets of miR398) play an important role in scavenging activity of ROS (results shown in Figure 8) [23]; SOD enzyme activity increased in both WT and lcd under PEG8000 (Figure 8A); Similarly, H2O2 and MDA contents increased in both WT and lcd under PEG8000 and it is notable that MDA content increased to a greater extent in lcd compared with WT (Figure 8B and 8C). The chemical compounds of mannitol, NaCl, and ABA were dissolved in MS medium, respectively. Wurzinger B, Mair A, Pfister B, Teige M. Cross-talk of calcium-dependent protein kinase and MAP kinase signaling. WT: wild type; #12, 25, 41: GmWRKY16 Arabidopsis transgenic lines of T3 generations. "Clearly, recently we have seen some very promising advances in terms of drought tolerances in crop plants," says Nguyen. ", "Open Access 'Chemistry' Journals allow the dissemination of knowledge at your finger tips without paying for the scientific content. 9:e27700. doi: 10.1104/pp.107.110981, Kim, S. Y. ( a, SF priming triggers a reduction in reactive oxygen species (ROS) levels. Transcription Factors (TFs) have a critical role in response to unregular and harsh environmental condition such as abiotic stresses through regulation of gene expression. The cotton WRKY transcription factor GhWRKY17 functions in drought and salt stress in transgenic Nicotiana benthamiana through ABA signaling and the modulation of reactive oxygen species production. The first Ministry of Health in Jordan was established in 1950. Similar resistant phenotypes to abiotic stress were also found in other WRKY TF genes in soybean. In the experiment, plants were re-watered on the day of wilting to allow the collection of leaf material after recovery. Different letters represent significant differences between the treatment means (, Stomatal closure response under drought. However, no significant differences in germination rate were found between WT and GmWRKY16 transgenic lines after 3 days under the treatments of 75, 100, and 150 mM NaCl, respectively (Figures 4C,D). Three independent biological experiments were performed. The purified PCR product was then inserted into the multiple cloning sites of the pLB vector (Tiangen Rapid DNA Ligation Kit, Beijing, China). Standardization of real-time PCR gene expression data from independent biological replicates. It is therefore possible that the decrease in tolerance and avoidance of drought stress was advantageous in the context of selection for increased competitive ability. The content of free proline in plants was determined following methods described previously (Bates et al., 1973). See this image and copyright information in PMC. Fast, scalable generation of high-quality protein multiple sequence alignments using Clustal Omega. Copyright: 2013 Shen et al. Physiol. Phylogenetic analysis was enforced by using the software of MEGA 7 (Kumar et al., 2016). The first day on which we observed that leaves had lost their turgidity was scored as wilting day. The WRKY TFs can be classified into three groups (I, II, and III) according to the number of WRKY domains and the Zn-finger motif type. SOD activities and malondialdehyde (MDA) content were measured according to Jiang et al. ", "It is important that students and researchers from all over the world can have easy access to relevant, high-standard and timely scientific information. Genome-wide identification of soybean WRKY transcription factors in response to salt stress. The WRKY TFs, dominating the genetic transcription, have become one of the largest TF families in plants (Eulgem et al., 2000; Rushton et al., 2010; Bencke-Malato et al., 2016; Goel et al., 2016; Meng et al., 2016; Yu Y. et al., 2016; Wu J. et al., 2017; Xiao et al., 2017; Bai et al., 2018; Finatto et al., 2018; Song et al., 2018; Xie et al., 2018). -, Fahad S., Bajwa A.A., Nazir U., Anjum S.A., Farooq A., Zohaib A., Sadia S., Nasim W., Adkins S., Saud S., et al. [19], ARF8 (target of miR167) prompts the elongation of hypocotyl and stamens during development, and regulates light signal transduction pathways. In a study on maize, K nutrition improved the drought tolerance, mainly due to improved cell membrane stability (Gnanasiri et al., 1991). (A) Expression patterns of genes responsive to salt stress. Aceituno FF, Moseyko N, Rhee SY, Gutirrez RA. Current Insights into the Molecular Mode of Action of Seaweed-Based Biostimulants and the Sustainability of Seaweeds as Raw Material Resources. Table S13: phenotypic data collected in this study. doi: 10.1002/2017EF000690. These results are consistent with an epistatic effect of the sos1 mutation over AVP1 overexpression in regard to salt tolerance ( Arabidopsis thaliana plants engineered to overexpress the vacuolar H+-pyrophosphatase AVP1 have enhanced tolerance to salinity and drought stress. Role of abscisic acid in plant stress tolerance. Stomatal density and length were quantified following the protocol described by Paccard et al. Jalalpour Z, Shabani L, Afghani L, Sharifi-Tehrani M. AMINI SA. Photosynthetic activity and duration of tolerated wilting were measured in the first experiment, whereas rosette area and leaf thickness were measured only in the second experiment (Supplementary Data Table S2). However, contrary phenotypes of drought tolerance by regulating ABI1/2 have also been reported over the past several years. Meanwhile, AtWRKY8 was induced with an expression level of over 12-fold greater under the NaCl treatment compared to that of the control (Figure 9A). On one hand, H2S directly regulates the expression of a series of drought responsive genes including DREB2A, DREB2B, RD29A and CBF4. (B) The determination of free proline content. BORKH., ENHANCES STOMATAL CLOSURE TO CONFER DROUGHT TOLERANCE IN TRANSGENIC ARABIDOPSIS AND APPLE Tan Y., Li M., Yang Y., Sun X., Wang N., Liang B., Ma F. Wu J, Chen J, Wang L, Wang S. Genome-Wide Investigation of WRKY Transcription Factors Involved in Terminal Drought Stress Response in Common Bean. ACTIN was used as an internal control in qRT-PCR. The papers published are of high quality after rigorous peer review and they are Indexed in: major international databases. The overall effect could lead to unexpected results such as in our example, where the expression of some target genes did not match that of their miRNAs. Liu J, Zhang F, Zhou J, Chen F, Wang B, Xie X. Lovell JT, Juenger TE, Michaels SD, et al. SF also positively affected stomatal behavior to support the tolerance to drought stress. Integrating circadian dynamics with physiological processes in plants. For this, plants were dried down a second time until wilting and re-watered after 3, 4, 5 or 6 d of wilting. Soybean WRKY-type transcription factor genes, GmWRKY13, GmWRKY21, and GmWRKY54, confer differential tolerance to abiotic stresses in transgenic Arabidopsis plants. doi: 10.1105/tpc.106.042705, Munns, R. (2002). In this study, ABI1 and ABI2 were upregulated by GmWRKY16, whereas ABI4 and ABI5 were downregulated in transgenic lines under drought stress (Figure 9C). Arch. Overexpression of TaWRKY146 increases drought tolerance through inducing stomatal closure in Arabidopsis thaliana. To examine the impact of plant size on the rate of soil water loss, we measured initial plant size and estimated the desiccation rate, defined as the rate of soil water loss per day over the 7 d following the water withdrawal in the second experiment of the dry-down experiment. These results established a significant correlation between drought stress and the production of both H2S transcripts and H2S emission. SF also positively affected stomatal behavior to support the tolerance to drought stress. (2017). (A) SOD activity was measured in WT and lcd plants treated with 0.2 g ml1 PEG8000 for 2 h. One SOD unit was the amount of enzyme required to inhibit photoreduction of nitro blue tetrazolium chIoride by 50% at 25C. These both sequences have the highly 5-bp conserved sequence of CCGAC that has the ability to transcription regulation in drought, salinity and temperature [128]. To identify the functional key genes, we used non-repetitive genes distinct in each functional category. Water loss of fresh leaves was weighed at treatment intervals set as 0, 0.5, 1, 1.5, 2, 3, 4, 5, and 6 h, respectively. The maize WRKY transcription factor ZmWRKY17 negatively regulates salt stress tolerance in transgenic Arabidopsis plants. Mitogen-Activated Protein Kinases (MAPKs) [36-38], Glycogen Synthase Kinase3 (GSK3) [40-41], S6 Kinase (S6K) [42], Calcium-Dependent Protein Kinases (CDPKs) [43-45] and most of SNF1-related kinases (SnRKs). Zhifang G., Loescher W. H. Expression of a celery man nose 6 phosphate reductase in Arabidopsis thaliana enhances salt tolerance and induces biosynthesis of both mannitol 125.Rivero R.M., Kojima M., Gepstein A. et al. These findings suggest that GmWRKY16 might play a role in multiple abiotic stresses. They provide easy access to the latest research on a wide variety of issues. 4B; Supplementary Data Table S9). 83, 265277. In contrast, a greater MDA accumulation of 0.019 mol.g-1 of fresh weight was found in WT compared to those of GmWRKY16 transgenic lines, with less than 0.013 mol.g-1 of fresh weight under salt stress (Figure 6C). (C) The determination of MDA content. In a previous study, we found that H2S interacts with ABA in the stomatal regulation of drought stress in Arabidopsis [5]. ", "Open access journals represent a major break-through in publishing. Since various stresses induce ABA synthesis, it is considered as a plant stress hormone [61, 62]. The 3-day-old seedlings from 1/2 MS medium were transferred to the plates of 1/2 MS containing ABA (0, 0.5, and 1.0 M). This is exactly what Open Access Journals provide and this is the reason why I support this endeavor. Consumption of H2O2 was measured as the decrease in absorbance at 470 nm. ", "Open access journals have become a fundamental tool for students, researchers, patients and the general public.
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