Abiotic stress

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    Environmental drivers and genomic architecture of trait differentiation in fire-adapted Banksia attenuata ecotypes
    Tianhua He, Byron B. Lamont, Neal J. Enright, Haylee M. D’Agui and William Stock
    J Integr Plant Biol 2019, 61 (4): 417-432.  
    DOI: 10.1111/jipb.12697
    Abstract (Browse 170)  |   Save
    Trait divergence between populations is considered an adaptive response to different environments, but to what extent this response is accompanied by genetic differentiation is less clear since it may be phenotypic plasticity. In this study, we analyzed phenotypic variation between two Banksia attenuata growth forms, lignotuberous (shrub) and epicormic resprouting (tree), in fire-prone environments to identify the environmental factors that have driven this phenotypic divergence. We linked genotype with phenotype and traced candidate genes using differential gene expression analysis. Fire intervals determined the phenotypic divergence between growth forms in B. attenuata. A genome-wide association study identified 69 single nucleotide polymorphisms, putatively associated with growth form, whereas no growth form- or phenotype-specific genotypes were identified. Genomic differentiation between the two growth forms was low (Fst = 0.024). Differential gene expression analysis identified 37 genes/transcripts that were differentially expressed in the two growth forms. A small heat-shock protein gene, associated with lignotuber presence, was differentially expressed in the two forms. We conclude that different fire regimes induce phenotypic polymorphism in B. attenuata, whereas phenotypic trait divergence involves the differential expression of a small fraction of genes that interact strongly with the disturbance regime. Thus, phenotypic plasticity among resprouters is the general strategy for surviving varying fire regimes.
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    ZmOST1 mediates abscisic acid regulation of guard cell ion channels and drought stress responses
    Qiqi Wu, Mei Wang, Jianlin Shen, Donghua Chen, Yu Zheng and Wei Zhang
    J Integr Plant Biol 2019, 61 (4): 478-491.  
    DOI: 10.1111/jipb.12714
    Abstract (Browse 321)  |   Save

    The phytohormone abscisic acid (ABA) is an important mediator in the drought response, participating in, among other processes, stomatal movements. In Arabidopsis thaliana, the serine/threonine protein kinase, OST1, regulates this response, but the function of its maize homolog has yet to be established. Here, we isolated ZmOST1 and show that its encoded protein indeed acts to regulate guard cell movement. ZmOST1 was ubiquitously expressed throughout the plant, being highly expressed in guard cells, and inducible both by exogenous ABA and water stress. Transient expression of a ZmOST1-GFP fusion protein, in maize mesophyll protoplasts, indicated its subcellular localization in the cytoplasm and nucleus. A Zmost1 loss-of-function mutant exhibited reduced sensitivity to ABA-activated slow anion channels in maize guard cells, and reduced drought tolerance. Constitutive expression of ZmOST1, in an A. thaliana ost1-1 mutant rescued the phenotype with respect both to the sensitivity of guard cell slow anion currents to ABA treatment and stomatal closure. Our findings indicate a positive regulatory role for ZmOST1 in guard cell ABA signaling and drought response in maize plants.

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    Endoplasmic reticulum stress-induced accumulation of VAMP721/722 requires CALRETICULIN 1 and CALRETICULIN 2 in Arabidopsis
    Soohong Kim, Yunjin Choi, Chian Kwon and Hye Sup Yun
    J Integr Plant Biol 2019, 61 (9): 974-980.  
    doi: 10.1111/jipb.12728
    Abstract (Browse 236)  |   Save

    Excessive demand for translation and protein folding in the endoplasmic reticulum (ER) can cause ER stress in plants. Here, we show that CALRETICULIN 1 (CRT1) and CRT2 are critical components in the accumulation of VESICLE‐ASSOCIATED MEMBRANE PROTEIN 721 (VAMP721) and VAMP722 during ER stress responses. We show that CRT2 interacts with VAMP722 and that CRT1/2 post‐translationally maintain elevated VAMP721/722 levels under ER stress. The greater growth inhibition in VAMP721/722‐deficient plants, induced by tunicamycin, suggests that plants under ER stress maintain physiological homeostasis, at least in part, by regulating VAMP721/722 levels, as VAMP721/722 are known to participate in various biological processes.

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    Basic-leucine zipper 17 and Hmg-CoA reductase degradation 3A are involved in salt acclimation memory in Arabidopsis
    Lin Tian, Yan Zhang, Erfang Kang, Huifang Ma, Huan Zhao, Ming Yuan, Lei Zhu and Ying Fu
    J Integr Plant Biol 2019, 61 (10): 1062-1084.  
    doi: 10.1111/jipb.12744
    Abstract (Browse 250)  |   Save
    Salt acclimation, which is induced by previous salt exposure, increases the resistance of plants to future exposure to salt stress. However, little is known about the underlying mechanism, particularly how plants store the “memory” of salt exposure. In this study, we established a system to study salt acclimation in Arabidopsis thaliana. Following treatment with a low concentration of salt, seedlings were allowed to recover to allow transitory salt responses to subside while maintaining the sustainable effects of salt acclimation. We performed transcriptome profiling analysis of these seedlings to identify genes related to salt acclimation memory. Notably, the expression of Basic‐leucine zipper 17 (bZIP17) and Hmg‐CoA reductase degradation 3A (HRD3A), which are important in the unfolded protein response (UPR) and endoplasmic reticulum‐associated degradation (ERAD), respectively, increased following treatment with a low concentration of salt and remained at stably high levels after the stimulus was removed, a treatment which improved plant tolerance to future high‐salinity challenge. Our findings suggest that the upregulated expression of important genes involved in the UPR and ERAD represents a “memory” of the history of salt exposure and enables more potent responses to future exposure to salt stress, providing new insights into the mechanisms underlying salt acclimation in plants.
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    Nitric oxide, γ-aminobutyric acid, and mannose pretreatment influence metabolic profiles in white clover under water stress
    Zhou Li, Bin Yong, Bizhen Cheng, Xing Wu, Yan Zhang, Xinquan Zhang and Yan Peng
    J Integr Plant Biol 2019, 61 (12): 1255-1273.  
    DOI: 10.1111/jipb.12770
    Abstract (Browse 440)  |   Save
    Nitric oxide (NO), γ‐aminobutyric acid (GABA), and mannose (MAS) could be important regulators of plant growth and adaptation to water stress. The application of sodium nitroprusside (SNP, a NO donor), GABA, and MAS improved plant growth under water‐sufficient conditions and effectively mitigated water stress damage to white clover. The metabonomic analysis showed that both SNP and GABA application resulted in a significant increase in myo‐inositol content; the accumulation of mannose was commonly regulated by SNP and MAS; GABA and MAS induced the accumulation of aspartic acid, quinic acid, trehalose, and glycerol under water deficit. In addition, citric acid was uniquely up‐regulated by SNP associated with tricarboxylic acid (TCA) cycle under water stress. GABA specially induced the accumulation of GABA, glycine, methionine, and aconitic acid related to GABA shunt, amino acids metabolism, and TCA cycle in response to water stress. MAS uniquely enhanced the accumulation of asparagine, galactose, and D‐pinitol in association with amino acids and sugars metabolism under water stress. SNP‐, GABA‐, and MAS‐induced changes of metabolic profiles and associated metabolic pathways could contribute to enhanced stress tolerance via involvement in the TCA cycle for energy supply, osmotic adjustment, antioxidant defense, and signal transduction for stress defense in white clover.
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    Induction of desiccation tolerance in desiccation sensitive Citrus limon seeds
    Alexandre Marques, Harm Nijveen, Charles Somi, Wilco Ligterink and Henk Hilhorst
    J Integr Plant Biol 2019, 61 (5): 624-638.  
    doi: 10.1111/jipb.12788
    Abstract (Browse 384)  |   Save
    Many economically important perennial species bear recalcitrant seeds, including tea, coffee, cocoa, mango, citrus, rubber, oil palm and coconut. Orthodox seeds can be dried almost completely without losing viability, but so-called recalcitrant seeds have a very limited storage life and die upon drying below a higher critical moisture content than orthodox seeds. As a result, the development of long-term storage methods for recalcitrant seeds is compromised. Lowering this critical moisture content would be very valuable since dry seed storage is the safest, most convenient and cheapest method for conserving plant genetic resources. Therefore, we have attempted to induce desiccation tolerance (DT) in the desiccation sensitive seeds of Citrus limon. We show that DT can be induced by paclobutrazol (an inhibitor of gibberellin biosynthesis) and we studied its associated transcriptome to delineate the molecular mechanisms underlying this induction of DT. Paclobutrazol not only interfered with gibberellin related gene expression but also caused extensive changes in expression of genes involved in the biosynthesis and signaling of other hormones. Paclobutrazol induced a transcriptomic switch encompassing suppression of biotic- and induction of abiotic responses. We hypothesize that this is the main driver of the induction of DT by paclobutrazol in C. limon seeds.
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    Genome-wide association study dissects the genetic bases of salt tolerance in maize seedlings
    Xi Luo, Bingcai Wang, Shan Gao, Fei Zhang, William Terzaghi and Mingqiu Dai
    J Integr Plant Biol 2019, 61 (6): 658-674.  
    doi: 10.1111/jipb.12797
    Abstract (Browse 1060)  |   Save
    Excess salinity is a natural stress that causes crop yield losses worldwide. The genetic bases of maize salt tolerance remain largely unknown. Here we investigated the survival rates of 445 maize natural accessions after salt treatments. A skewed distribution of the salt-tolerant phenotypes was observed in this population. Genome-wide association studies (GWAS) revealed 57 loci significantly associated with salt tolerance. Forty-nine candidate genes were detected from these loci. About 10% of these genes were co-localized with loci from QTL mapping. Forty four percent of the candidate genes were involved in stress responses, ABA signaling, stomata division, DNA binding/transcription regulation and auxin signaling, suggesting that they are key genetic mechanisms of maize salt tolerance. Transgenic studies showed that two genes, the salt-tolerance-associated-gene 4 (SAG4, GRMZM2G077295) and SAG6 (GRMZM2G106056), which encode a protein transport protein and the double-strand break repair protein MRE11, respectively, had positive roles in plant salt tolerance, and their salt-tolerant haplotypes were revealed. The genes we identified in this study provide a list of candidate targets for further study of maize salt tolerance, and of genetic markers and materials that may be used for breeding salt-tolerance in maize.
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    The involvement of wheat U‐box E3 ubiquitin ligase TaPUB1 in salt stress tolerance
    Wenlong Wang, Wenqiang Wang, Yunzhen Wu, Qinxue Li, Guangqiang Zhang, Ruirui Shi, Junjiao Yang, Yong Wang and Wei Wang
    J Integr Plant Biol 2020, 62 (5): 631-651.  
    DOI: 10.1111/jipb.12842
    Abstract (Browse 378)  |   Save

    U‐box E3 ubiquitin ligases play important roles in the ubiquitin/26S proteasome machinery and in abiotic stress responses. TaPUB1‐overexpressing wheat (Triticum aestivum L.) were generated to evaluate its function in salt tolerance. These plants had more salt stress tolerance during seedling and flowering stages, whereas the TaPUB1‐RNA interference (RNAi)‐mediated knock‐down transgenic wheat showed more salt stress sensitivity than the wild type (WT). TaPUB1 overexpression upregulated the expression of genes related to ion channels and increased the net root Na+ efflux, but decreased the net K+ efflux and H+ influx, thereby maintaining a low cytosolic Na+/K+ ratio, compared with the WT. However, RNAi‐mediated knock‐down plants showed the opposite response to salt stress. TaPUB1 could induce the expression of some genes that improved the antioxidant capacity of plants under salt stress. TaPUB1 also interacted with TaMP (Triticum aestivum α‐mannosidase protein), a regulator playing an important role in salt response in yeast and in plants. Thus, low cytosolic Na+/K+ ratios and better antioxidant enzyme activities could be maintained in wheat with overexpression of TaPUB1 under salt stress. Therefore, we conclude that the U‐box E3 ubiquitin ligase TaPUB1 positively regulates salt stress tolerance in wheat.

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    Screening of abiotic stress-responsive cotton genes using a cotton full-length cDNA overexpressing Arabidopsis library
    Shengting Li, Hao Chen, Zhi Hou, Yu Li, Cuiling Yang, DaoJie Wang and Chun‐Peng Song
    J Integr Plant Biol 2020, 62 (7): 998-1016.  
    DOI: 10.1111/jipb.12861
    Abstract (Browse 470)  |   Save

    Cotton (Gossypium hirsutum L.) is a major crop and the main source of natural fiber worldwide. Because various abiotic and biotic stresses strongly influence cotton fiber yield and quality, improved stress resistance of this crop plant is urgently needed. In this study, we used Gateway technology to construct a normalized full‐length cDNA overexpressing (FOX) library from upland cotton cultivar ZM12 under various stress conditions. The library was transformed into Arabidopsis to produce a cotton‐FOX‐Arabidopsis library. Screening of this library yielded 6,830 transgenic Arabidopsis lines, of which 757 were selected for sequencing to ultimately obtain 659 cotton ESTs. GO and KEGG analyses mapped most of the cotton ESTs to plant biological process, cellular component, and molecular function categories. Next, 156 potential stress‐responsive cotton genes were identified from the cotton‐FOX‐Arabidopsis library under drought, salt, ABA, and other stress conditions. Four stress‐related genes identified from the library, designated as GhCAS , GhAPX , GhSDH , and GhPOD , were cloned from cotton complementary DNA, and their expression patterns under stress were analyzed. Phenotypic experiments indicated that overexpression of these cotton genes in Arabidopsis affected the response to abiotic stress. The method developed in this study lays a foundation for high‐throughput cloning and rapid identification of cotton functional genes.

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    S -acylation of CBL10/SCaBP8 by PAT10 is crucial for its tonoplast association and function in salt tolerance
    Sen Chai, Fu-Rong Ge, Yan Zhang and Sha Li
    J Integr Plant Biol 2020, 62 (6): 718-722.  
    doi: 10.1111/jipb.12864
    Abstract (Browse 305)  |   Save

    Crop yield is sensitive to salt stresses, for which Calcineurin B‐like proteins (CBLs) are major response factors. This study shows that Arabidopsis CBL10, through protein S ‐acylation by protein S ‐acyl transferase10, targets to the vacuolar membrane to confer salt tolerance.

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    Reactive oxygen species regulate auxin levels to mediate adventitious root induction in Arabidopsis hypocotyl cuttings
    Aixia Huang, Yongshun Wang, Yangyang Liu, Guodong Wang and Xiaoping She
    J Integr Plant Biol 2020, 62 (7): 912-926.  
    DOI: 10.1111/jipb.12870
    Abstract (Browse 373)  |   Save

    Adventitious root (AR) formation from leafy stem cuttings is critical for breeding of many forest and horticultural species. In addition to the plant hormone auxin, wound‐induced signaling caused by the cutting excision is also essential for AR initiation. Here we found that reactive oxygen species (ROS) are rapidly generated at the excision site as a wound‐induced signal and propagated throughout the hypocotyl cutting after excision of the Arabidopsis (Arabidopsis thaliana ) primary root. ROS propagation was not observed in the presence of an NADPH oxidase inhibitor (diphenylene iodonium chloride) or in a knockout mutant of the NADPH oxidase gene respiratory burst oxidase homolog protein D (RBOHD ). Respiratory burst oxidase homolog protein D was specifically upregulated in hypocotyl cuttings at 0.5 h post excision (hpe). Together, these data suggest that RBOHD mediates ROS propagation in hypocotyl cuttings. We also found that auxin levels increased significantly in the shoot apex at 5 hpe and at the base of the cutting at 6 hpe; these effects were blocked by treatment with ROS scavengers. Consistent with this, transcript levels of auxin biosynthesis and polar‐transport genes generally increased between 1 to 6 hpe. Collectively, our results suggest that wound‐induced ROS participate in AR induction through regulation of auxin biosynthesis and transport.

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    Polyunsaturated linolenoyl‐CoA modulates ERF‐VII‐mediated hypoxia signaling in Arabidopsis
    Ying Zhou, Wei-Juan Tan, Li-Juan Xie, Hua Qi, Yi-Cong Yang, Li-Ping Huang, Yong-Xia Lai, Yi-Fang Tan, De-Mian Zhou, Lu-Jun Yu, Qin-Fang Chen, Mee-Len Chye and Shi Xiao
    J Integr Plant Biol 2020, 62 (3): 330-348.  
    DOI: 10.1111/jipb.12875
    Abstract (Browse 386)  |   Save

    In plants, submergence from flooding causes hypoxia, which impairs energy production and affects plant growth, productivity, and survival. In Arabidopsis, hypoxia induces nuclear localization of the group VII ethylene‐responsive transcription factor RELATED TO AP2.12 (RAP2.12), following its dissociation from the plasma membrane‐anchored ACYL‐COA BINDING PROTEIN1 (ACBP1) and ACBP2. Here, we show that polyunsaturated linolenoyl‐CoA (18:3‐CoA) regulates RAP2.12 release from the plasma membrane. Submergence caused a significant increase in 18:3‐CoA, but a significant decrease in 18:0‐, 18:1‐, and 18:2‐CoA. Application of 18:3‐CoA promoted nuclear accumulation of the green fluorescent protein (GFP) fusions RAP2.12‐GFP, HYPOXIA‐RESPONSIVE ERF1‐GFP, and RAP2.3‐GFP, and enhanced transcript levels of hypoxia‐responsive genes. Plants with decreased ACBP1 and ACBP2 (acbp1 ACBP2‐RNAi, produced by ACBP2 RNA interference in the acbp1 mutant) had reduced tolerance to hypoxia and impaired 18:3‐CoA‐induced expression of hypoxia‐related genes. In knockout mutants and overexpression lines of LONG‐CHAIN ACYL‐COA SYNTHASE2 (LACS2) and FATTY ACID DESATURASE 3 (FAD3), the acyl‐CoA pool size and 18:3‐CoA levels were closely related to ERF‐VII‐mediated signaling and hypoxia tolerance. These findings demonstrate that polyunsaturation of long‐chain acyl‐CoAs functions as important mechanism in the regulation of plant hypoxia signaling, by modulating ACBP–ERF‐VII dynamics.

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    Overexpression of CmSOS1 confers waterlogging tolerance in Chrysanthemum
    Lijun Wang, Jiaojiao Gao, Zixin Zhang, Weimiao Liu, Peilei Cheng, Wenting Mu, Tong Su, Sumei Chen, Fadi Chen and Jiafu Jiang
    J Integr Plant Biol 2020, 62 (8): 1059-1064.  
    doi: 10.1111/jipb.12889
    Abstract (Browse 649)  |   Save

    The Na+/H+ antiporter SOS1 enhances the salinity tolerance of a number of plant species, but its involvement in the response to hypoxia is less well known. We presented chrysanthemum homologs CmSOS1 and CmRCD1 coordinately mediate waterlogging tolerance by maintaining membrane integrity and minimizing the level of reactive oxygen species.

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    New insights into gibberellin signaling in regulating flowering in Arabidopsis
    Shengjie Bao, Changmei Hua, Lisha Shen and Hao Yu
    J Integr Plant Biol 2020, 62 (1): 118-131.  
    doi: 10.1111/jipb.12892
    Abstract (Browse 506)  |   Save

    In angiosperms, floral transition is a key developmental transition from the vegetative to reproductive growth, and requires precise regulation to maximize the reproductive success. A complex regulatory network governs this transition through integrating flowering pathways in response to multiple exogenous and endogenous cues. Phytohormones are essential for proper plant developmental regulation and have been extensively studied for their involvement in the floral transition. Among various phytohormones, gibberellin (GA) plays a major role in affecting flowering in the model plant Arabidopsis thaliana. The GA pathway interact with other flowering genetic pathways and phytohormone signaling pathways through either DELLA proteins or mediating GA homeostasis. In this review, we summarize the recent advances in understanding the mechanisms of DELLA‐mediated GA pathway in flowering time control in Arabidopsis, and discuss its possible link with other phytohormone pathways during the floral transition.

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    The roles of endomembrane trafficking in plant abiotic stress responses
    Xiangfeng Wang, Min Xu, Caiji Gao, Yonglun Zeng, Yong Cui, Wenjin Shen and Liwen Jiang
    J Integr Plant Biol 2020, 62 (1): 55-69.  
    doi: 10.1111/jipb.12895
    Abstract (Browse 368)  |   Save

    Endomembrane trafficking is a fundamental cellular process in all eukaryotic cells and its regulatory mechanisms have been extensively studied. In plants, the endomembrane trafficking system needs to be constantly adjusted to adapt to the ever‐changing environment. Evidence has accumulated supporting the idea that endomembrane trafficking is tightly linked to stress signaling pathways to meet the demands of rapid changes in cellular processes and to ensure the correct delivery of stress‐related cargo molecules. However, the underlying mechanisms remain unknown. In this review, we summarize the recent findings on the functional roles of both secretory trafficking and endocytic trafficking in different types of abiotic stresses. We also highlight and discuss the unique properties of specific regulatory molecules beyond their conventional functions in endosomal trafficking during plant growth under stress conditions.

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    Epigenetic regulation in plant abiotic stress responses
    Ya-Nan Chang, Chen Zhu, Jing Jiang, Huiming Zhang, Jian-Kang Zhu and Cheng-Guo Duan
    J Integr Plant Biol 2020, 62 (5): 563-580.  
    doi: 10.1111/jipb.12901
    Abstract (Browse 903)  |   Save

    In eukaryotic cells, gene expression is greatly influenced by the dynamic chromatin environment. Epigenetic mechanisms, including covalent modifications to DNA and histone tails and the accessibility of chromatin, create various chromatin states for stress‐responsive gene expression that is important for adaptation to harsh environmental conditions. Recent studies have revealed that many epigenetic factors participate in abiotic stress responses, and various chromatin modifications are changed when plants are exposed to stressful environments. In this review, we summarize recent progress on the cross‐talk between abiotic stress response pathways and epigenetic regulatory pathways in plants. Our review focuses on epigenetic regulation of plant responses to extreme temperatures, drought, salinity, the stress hormone abscisic acid, nutrient limitations and ultraviolet stress, and on epigenetic mechanisms of stress memory.

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    Trehalose‐6‐phosphate phosphatase E modulates ABA‐controlled root growth and stomatal movement in Arabidopsis
    Wenjing Wang, Qingbin Chen, Shouming Xu, Wen-Cheng Liu, Xiaohong Zhu, and Chun-Peng Song
    J Integr Plant Biol 2020, 62 (10): 1518-1534.  
    doi: 10.1111/jipb.12925
    Abstract (Browse 517)  |   Save

    Trehalose plays important roles in plant growth and stress responses and is synthesized from trehalose‐6‐phosphate by trehalose‐6‐phosphate phosphatase (TPP). Here, we show that trehalose and abscisic acid (ABA) have synergistic effects on root growth and stomatal closure. The Arabidopsis thaliana genome contains ten genes encoding TPPs and the expression level of one, TPPE, and trehalose contents increased in response to ABA. In the presence of ABA, the ABA‐responsive transcription factor ABA RESPONSE ELEMENT BINDING FACTOR2 (ABF2) directly binds to the TPPE promoter to activate its expression. Genetic analysis revealed that TPPE acts downstream of ABF2, which is supported by the findings that TPPE expression and trehalose content are reduced in the abf2 mutant and that a mutation in TPPE abolished the ABA‐sensitive root elongation phenotype of 35S:ABF2 plants. Reactive oxygen species (ROS) accumulation in response to ABA failed to occur in tppe mutant plants, suggesting that TPPE is involved in ABA‐controlled root elongation and stomatal movement by inducing ROS accumulation. This study uncovers a new branch of the ABA signaling pathway and provides a molecular basis for the role of trehalose in plant responses to abiotic stress.

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    BTB/TAZ protein MdBT2 integrates multiple hormonal and environmental signals to regulate anthocyanin biosynthesis in apple
    Jian-Ping An, Xiao-Fei Wang and Yu-Jin Hao
    J Integr Plant Biol 2020, 62 (11): 1643-1646.  
    doi: 10.1111/jipb.12940
    Abstract (Browse 363)  |   Save

    BT2 is a BTB/TAZ domain protein with key roles in multiple stress responses and the plant development of Arabidopsis (Figueroa et al. 2005; Ren et al. 2007; Mandadi et al. 2009). Recent studies have demonstrated that apple MdBT2 functions as a negative regulator in diverse hormonal and environmental signal‐induced anthocyanin biosynthesis, suggesting that MdBT2 integrates stress signals and anthocyanin biosynthesis.

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    ZmSRL5 is involved in drought tolerance by maintaining cuticular wax structure in maize
    Zhenyuan Pan, Min Liu, Hailiang Zhao, Zengdong Tan, Kun Liang, Qin Sun, Dianming Gong, Haijun He, Wenqi Zhou and Fazhan Qiu
    J Integr Plant Biol 2020, 62 (12): 1895-1909.  
    DOI: 10.1111/jipb.12982
    Abstract (Browse 457)  |   Save

    Cuticular wax is a natural barrier on terrestrial plant organs, which protects plants from damages caused by a variety of stresses. Here, we report the identification and functional characterization of a cuticular‐wax‐related gene, Zea mays L. SEMI‐ROLLED LEAF 5 (ZmSRL5). The loss‐of‐function mutant srl5, which was created by a 3,745 bp insertion in the first intron that led to the premature transcript, exhibited abnormal wax crystal morphology and distribution, which, in turn, caused the pleiotropic phenotypes including increased chlorophyll leaching and water loss rate, decreased leaf temperature, sensitivity to drought, as well as semi‐rolled mature leaves. However, total wax amounts showed no significant difference between wild type and semi‐rolled leaf5 (srl5) mutant. The phenotype of srl5 was confirmed through the generation of two allelic mutants using CRISPR/Cas9. ZmSRL5 encodes a CASPARIAN‐STRIP‐MEMBRANE‐DOMAIN‐LIKE (CASPL) protein located in plasma membrane, and highly expressed in developing leaves. Further analysis showed that the expressions of the most wax related genes were not affected or slightly altered in srl5. This study, thus, primarily uncovers that ZmSRL5 is required for the structure formation of the cuticular wax and could increase the drought tolerance by maintaining the proper cuticular wax structure in maize.

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    Melatonin: A master regulator of plant development and stress responses
    Chengliang Sun, Lijuan Liu, Luxuan Wang, Baohai Li, Chongwei Jin and Xianyong Lin
    J Integr Plant Biol 2021, 63 (1): 126-145.  
    doi: 10.1111/jipb.12993
    Abstract (Browse 540)  |   Save
    Melatonin is a pleiotropic molecule with multiple functions in plants. Since the discovery of melatonin in plants, numerous studies have provided insight into the biosynthesis, catabolism, and physiological and biochemical functions of this important molecule. Here, we describe the biosynthesis of melatonin from tryptophan, as well as its various degradation pathways in plants. The identification of a putative melatonin receptor in plants has led to the hypothesis that melatonin is a hormone involved in regulating plant growth, aerial organ development, root morphology, and the floral transition. The universal antioxidant activity of melatonin and its role in preserving chlorophyll might explain its anti‐senescence capacity in aging leaves. An impressive amount of research has focused on the role of melatonin in modulating postharvest fruit ripening by regulating the expression of ethylene‐related genes. Recent evidence also indicated that melatonin functions in the plant's response to biotic stress, cooperating with other phytohormones and well‐known molecules such as reactive oxygen species and nitric oxide. Finally, great progress has been made towards understanding how melatonin alleviates the effects of various abiotic stresses, including salt, drought, extreme temperature, and heavy metal stress. Given its diverse roles, we propose that melatonin is a master regulator in plants.
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    Betula platyphylla BpHOX2 transcription factor binds to different cis-acting elements and confers osmotic tolerance
    Zilong Tan , Xuejing Wen and Yucheng Wang
    J Integr Plant Biol 2020, 62 (11): 1762-1779.  
    DOI: 10.1111/jipb.12994
    Abstract (Browse 392)  |   Save

    The homeodomain‐leucine zipper (HD‐Zip) proteins play crucial roles in plant developmental and environmental responses. However, how they mediate gene expression to facilitate abiotic stress tolerance remains unknown. In the present study, we characterized BpHOX2 (encoding a HD‐Zip I family protein) from birch (Betula platyphylla). BpHOX2 is predominately expressed in mature stems and leaves, but expressed at a low level in apical buds and roots, suggesting that it has tissue‐specific characteristics. BpHOX2 expression was highly induced by osmotic and salt, but only slightly induced by abscisic acid. Overexpression of BpHOX2 markedly improved osmotic tolerance, while knockdown of BpHOX2 increased sensitivity to osmotic stress. BpHOX2 could induce the expression of pyrroline‐5‐carboxylate synthase, peroxidase, and superoxide dismutase genes to improve proline levels and the reactive oxygen species scavenging capability. Chromatin immunoprecipitation sequencing combined with RNA sequencing showed that BpHOX2 could bind to at least four cis‐acting elements, including dehydration‐responsive element “RCCGAC”, Myb‐p binding box “CCWACC,” and two novel cis‐acting elements with the sequences of “AAGAAG” and “TACGTG” (termed HBS1 and HBS2, respectively) to regulate gene expression. Our results suggested that BpHOX2 is a transcription factor that binds to different cis‐acting elements to regulate gene expression, ultimately improving osmotic tolerance in birch.

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    N4‐methylcytidine ribosomal RNA methylation in chloroplasts is crucial for chloroplast function, development, and abscisic acid response in Arabidopsis
    Le Nguyen Tieu Ngoc, Su Jung Park, Trinh Thi Huong, Kwang Ho Lee and Hunseung Kang
    J Integr Plant Biol 2021, 63 (3): 570-582.  
    doi: 10.1111/jipb.13009
    Abstract (Browse 309)  |   Save
    Although the essential role of messenger RNA methylation in the nucleus is increasingly understood, the nature of ribosomal RNA (rRNA) methyltransferases and the role of rRNA methylation in chloroplasts remain largely unknown. A recent study revealed that CMAL (for Chloroplast mr aW‐ Like) is a chloroplast‐localized rRNA methyltransferase that is responsible for N4‐methylcytidine (m4C) in 16S chloroplast rRNA in Arabidopsis thaliana. In this study, we further examined the role of CMAL in chloroplast biogenesis and function, development, and hormone response. The cmal mutant showed reduced chlorophyll biosynthesis, photosynthetic activity, and growth‐defect phenotypes, including severely stunted stems, fewer siliques, and lower seed yield. The cmal mutant was hypersensitive to chloroplast translation inhibitors, such as lincomycin and erythromycin, indicating that the m4C‐methylation defect in the 16S rRNA leads to a reduced translational activity in chloroplasts. Importantly, the stunted stem of the cmal mutant was partially rescued by exogenous gibberellic acid or auxin. The cmal mutant grew poorer than wild type, whereas the CMAL‐overexpressing transgenic Arabidopsis plants grew better than wild type in the presence of abscisic acid. Altogether, these results indicate that CMAL is an indispensable rRNA methyltransferase in chloroplasts and is crucial for chloroplast biogenesis and function, photosynthesis, and hormone response during plant growth and development.
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    The Arabidopsis phosphatase PP2C49 negatively regulates salt tolerance through inhibition of AtHKT1;1
    Moli Chu , Pengwang Chen, Sufang Meng, Peng Xu and Wenzhi Lan
    J Integr Plant Biol 2021, 63 (3): 528-542.  
    doi: 10.1111/jipb.13008
    Abstract (Browse 522)  |   Save
    Type 2C protein phosphatases (PP2Cs) are the largest protein phosphatase family. PP2Cs dephosphorylate substrates for signaling in Arabidopsis, but the functions of most PP2Cs remain unknown. Here, we characterized PP2C49 (AT3G62260, a Group G PP2C), which regulates Na+ distribution under salt stress and is localized to the cytoplasm and nucleus. PP2C49 was highly expressed in root vascular tissues and its disruption enhanced plant tolerance to salt stress. Compared with wild type, the pp2c49 mutant contained more Na+ in roots but less Na+ in shoots and xylem sap, suggesting that PP2C49 regulates shoot Na+ extrusion. Reciprocal grafting revealed a root‐based mechanism underlying the salt tolerance of pp2c49. Systemic Na+ distribution largely depends on AtHKT1;1 and loss of function of AtHKT1;1 in the pp2c49 background overrode the salt tolerance of pp2c49, resulting in salt sensitivity. Furthermore, compared with plants overexpressing PP2C49 in the wild‐type background, plants overexpressing PP2C49 in the athtk1;1 mutant background were sensitive to salt, like the athtk1;1 mutants. Moreover, protein–protein interaction and two‐voltage clamping assays demonstrated that PP2C49 physically interacts with AtHKT1;1 and inhibits the Na+ permeability of AtHKT1;1. This study reveals that PP2C49 negatively regulates AtHKT1;1 activity and thus determines systemic Na+ allocation during salt stress.
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    Suppression of DRR1 results in the accumulation of insoluble ubiquitinated proteins, which impairs drought stress tolerance
    Seong Gwan Yu, Na Hyun Cho, Jong Hum Kim, Tae Rin Oh and Woo Taek Kim
    J Integr Plant Biol 2021, 63 (3): 431-437.  
    doi: 10.1111/jipb.13014
    Abstract (Browse 848)  |   Save
    Drought stress has detrimental effects on plants. Although the abscisic acid (ABA)‐mediated drought response is well established, defensive mechanisms to cope with dehydration‐induced proteotoxicity have been rarely studied. DRR1 was identified as an Arabidopsis drought‐induced gene encoding an ER‐localized RING‐type E3 Ub ligase. Suppression of DRR1 markedly reduced tolerance to drought and proteotoxic stress without altering ABA‐mediated germination and stomatal movement. Proteotoxicity‐ and dehydration‐induced insoluble ubiquitinated protein accumulation was more obvious in DRR1 loss‐of‐function plants than in wild‐type plants. These results suggest that DRR1 is involved in an ABA‐independent drought stress response possibly through the mitigation of dehydration‐induced proteotoxic stress.
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    RING finger protein RGLG1 and RGLG2 negatively modulate MAPKKK18 mediated drought stress tolerance in Arabidopsis
    Jiayi Yu, Lu Kang, Yuanyuan Li, Changai Wu, Chengchao Zheng, Pei Liu and Jinguang Huang
    J Integr Plant Biol 2021, 63 (3): 484-493.  
    doi: 10.1111/jipb.13019
    Abstract (Browse 540)  |   Save
    Mitogen activated protein kinase kinase kinase 18 (MAPKKK18) mediated signaling cascade plays important roles in Arabidopsis drought stress tolerance. However, the post‐translational modulation patterns of MAPKKK18 are not characterized. In this study, we found that the protein level of MAPKKK18 was tightly controlled by the 26S proteasome. Ubiquitin ligases RGLG1 and RGLG2 ubiquitinated MAPKKK18 at lysine residue K32 and K154, and promoted its degradation. Deletion of RGLG1 and RGLG2 stabilized MAPKKK18 and further enhanced the drought stress tolerance of MAPKKK18‐overexpression plants. Our data demonstrate that RGLG1 and RGLG2 negatively regulate MAPKKK18‐mediated drought stress tolerance in Arabidopsis.
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    Hydrogen sulfide, a signaling molecule in plant stress responses
    Jing Zhang, Mingjian Zhou, Heng Zhou, Didi Zhao, Cecilia Gotor, Luis C. Romero, Jie Shen, Zhenglin Ge, Zhirong Zhang, Wenbiao Shen, Xingxing Yuan and Yanjie Xie
    J Integr Plant Biol 2021, 63 (1): 146-160.  
    doi: 10.1111/jipb.13022
    Abstract (Browse 489)  |   Save
    Gaseous molecules, such as hydrogen sulfide (H2S) and nitric oxide (NO), are crucial players in cellular and (patho)physiological processes in biological systems. The biological functions of these gaseous molecules, which were first discovered and identified as gasotransmitters in animals, have received unprecedented attention from plant scientists in recent decades. Researchers have arrived at the consensus that H2S is synthesized endogenously and serves as a signaling molecule throughout the plant life cycle. However, the mechanisms of H2S action in redox biology is still largely unexplored. This review highlights what we currently know about the characteristics and biosynthesis of H2S in plants. Additionally, we summarize the role of H2S in plant resistance to abiotic stress. Moreover, we propose and discuss possible redox‐dependent mechanisms by which H2S regulates plant physiology.
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    ANNEXIN 8 negatively regulates RPW8.1‐mediated cell death and disease resistance in Arabidopsis
    Zhi‐Xue Zhao, Yong‐Ju Xu, Yang Lei, Qin Li, Ji‐Qun Zhao, Yan Li, Jing Fan, Shunyuan Xiao and Wen‐Ming Wang
    J Integr Plant Biol 2021, 63 (2): 378-392.  
    doi: 10.1111/jipb.13025
    Abstract (Browse 370)  |   Save
    Study on the regulation of broad‐spectrum resistance is an active area in plant biology. RESISTANCE TO POWDERY MILDEW 8.1 (RPW8.1) is one of a few broad‐spectrum resistance genes triggering the hypersensitive response (HR) to restrict multiple pathogenic infections. To address the question how RPW8.1 signaling is regulated, we performed a genetic screen and tried to identify mutations enhancing RPW8.1‐mediated HR. Here, we provided evidence to connect an annexin protein with RPW8.1‐mediated resistance in Arabidopsis against powdery mildew. We isolated and characterized Arabidopsis b7‐6 mutant. A point mutation in b7‐6 at the At5g12380 locus resulted in an amino acid substitution in ANNEXIN 8 (AtANN8). Loss‐of‐function or RNA‐silencing of AtANN8 led to enhanced expression of RPW8.1, RPW8.1‐dependent necrotic lesions in leaves, and defense against powdery mildew. Conversely, over‐expression of AtANN8 compromised RPW8.1‐mediated disease resistance and cell death. Interestingly, the mutation in AtANN8 enhanced RPW8.1‐triggered H2O2. In addition, mutation in AtANN8 led to hypersensitivity to salt stress. Together, our data indicate that AtANN8 is involved in multiple stress signaling pathways and negatively regulates RPW8.1‐mediated resistance against powdery mildew and cell death, thus linking ANNEXIN's function with plant immunity.
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    Comparative transcriptomics reveals hidden issues in the plant response to arthropod herbivores
    M. Estrella Santamaria, Alejandro Garcia, Ana Arnaiz, Irene Rosa‐Diaz, Gara Romero‐Hernandez, IsabelDiaz and Manuel Martinez
    J Integr Plant Biol 2021, 63 (2): 312-326.  
    doi: 10.1111/jipb.13026
    Abstract (Browse 313)  |   Save
    Plants experience different abiotic/biotic stresses, which trigger their molecular machinery to cope with them. Besides general mechanisms prompted by many stresses, specific mechanisms have been introduced to optimize the response to individual threats. However, these key mechanisms are difficult to identify. Here, we introduce an in‐depth species‐specific transcriptomic analysis and conduct an extensive meta‐analysis of the responses to related species to gain more knowledge about plant responses. The spider mite Tetranychus urticae was used as the individual species, several arthropod herbivores as the related species for meta‐analysis, and Arabidopsis thaliana plants as the common host. The analysis of the transcriptomic data showed typical common responses to herbivory, such as jasmonate signaling or glucosinolate biosynthesis. Also, a specific set of genes likely involved in the particularities of the Arabidopsis‐spider mite interaction was discovered. The new findings have determined a prominent role in this interaction of the jasmonate‐induced pathways leading to the biosynthesis of anthocyanins and tocopherols. Therefore, tandem individual/general transcriptomic profiling has been revealed as an effective method to identify novel relevant processes and specificities in the plant response to environmental stresses.
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    Ethylene signaling in rice and Arabidopsis: New regulators and mechanisms
    He Zhao, Cui-Cui Yin, Biao Ma, Shou-Yi Chen and Jin-Song Zhang
    J Integr Plant Biol 2021, 63 (1): 102-125.  
    doi: 10.1111/jipb.13028
    Abstract (Browse 460)  |   Save
    Ethylene is a gaseous hormone which plays important roles in both plant growth and development and stress responses. Based on studies in the dicot model plant species Arabidopsis, a linear ethylene signaling pathway has been established, according to which ethylene is perceived by ethylene receptors and transduced through CONSTITUTIVE TRIPLE RESPONSE 1 (CTR1) and ETHYLENE‐INSENSITIVE 2 (EIN2) to activate transcriptional reprogramming. In addition to this canonical signaling pathway, an alternative ethylene receptor‐mediated phosphor‐relay pathway has also been proposed to participate in ethylene signaling. In contrast to Arabidopsis, rice, a monocot, grows in semiaquatic environments and has a distinct plant structure. Several novel regulators and/or mechanisms of the rice ethylene signaling pathway have recently been identified, indicating that the ethylene signaling pathway in rice has its own unique features. In this review, we summarize the latest progress and compare the conserved and divergent aspects of the ethylene signaling pathway between Arabidopsis and rice. The crosstalk between ethylene and other plant hormones is also reviewed. Finally, we discuss how ethylene regulates plant growth, stress responses and agronomic traits. These analyses should help expand our knowledge of the ethylene signaling mechanism and could further be applied for agricultural purposes.
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    Abscisic acid receptors maintain abscisic acid homeostasis by modulating UGT71C5 glycosylation activity
    Yanlin Ma, Jing Cao, Qiaoqiao Chen, Jiahan He, Zhibin Liu, Jianmei Wang, Xufeng Li and Yi Yang
    J Integr Plant Biol 2021, 63 (3): 543-552.  
    doi: 10.1111/jipb.13030
    Abstract (Browse 387)  |   Save
    Uridine diphosphate‐glucosyltransferases (UGTs) maintain abscisic acid (ABA) homeostasis in Arabidopsis thaliana by converting ABA to abscisic acid‐glucose ester (ABA‐GE). UGT71C5 plays an important role in the generation of ABA‐GE. Abscisic acid receptors are crucial upstream components of the ABA signaling pathway, but how UGTs and ABA receptors function together to modulate ABA levels is unknown. Here, we demonstrated that the ABA receptors RCAR12/13 and UGT71C5 maintain ABA homeostasis in Arabidopsis following rehydration under drought stress. Biochemical analyses show that UGT71C5 directly interacted with RCAR8/12/13 in yeast cells, and the interactions between UGT71C5 and RCAR12/13 were enhanced by ABA treatment. Enzyme activity analysis showed that ABA‐GE contents were significantly elevated in the presence of RCAR12 or RCAR13, suggesting that these ABA receptors enhance the activity of UGT71C5. Determination of the content of ABA and ABA‐GE in Arabidopsis following rehydration under drought stress revealed that ABA‐GE contents were significantly higher in Arabidopsis plants overexpressing RCAR12 and RCAR13 than in non‐transformed plants and plants overexpressing RCAR11 following rehydration under drought stress. These observations suggest that RCAR12 and RCAR13 enhance the activity of UGT71C5 to glycosylate excess ABA into ABA‐GE following rehydration under drought stress, representing a rapid mechanism for regulating plant growth and development.
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    Roles of DEMETER in regulating DNA methylation in vegetative tissues and pathogen resistance
    Wenjie Zeng, Huan Huang, Xueqiang Lin, Chen Zhu, Ken‐ichi Kosami, Chaofeng Huang, Huiming Zhang, Cheng‐Guo Duan, Jian‐Kang Zhu and Daisuke Miki
    J Integr Plant Biol 2021, 63 (4): 691-706.  
    doi: 10.1111/jipb.13037
    Abstract (Browse 301)  |   Save
    DNA methylation is an epigenetic mark important for genome stability and gene expression. In Arabidopsis thaliana, the 5‐methylcytosine DNA glycosylase/demethylase DEMETER (DME) controls active DNA demethylation during the reproductive stage; however, the lethality of loss‐of‐function dme mutations has made it difficult to assess DME function in vegetative tissues. Here, we edited DME using clustered regularly interspaced short palindromic repeats (CRISPR) /CRISPR‐associated protein 9 and created three weak dme mutants that produced a few viable seeds. We also performed central cell‐specific complementation in a strong dme mutant and combined this line with mutations in the other three Arabidopsis demethylase genes to generate the dme ros1 dml2 dml3 (drdd) quadruple mutant. A DNA methylome analysis showed that DME is required for DNA demethylation at hundreds of genomic regions in vegetative tissues. A transcriptome analysis of the drdd mutant revealed that DME and the other three demethylases are important for plant responses to biotic and abiotic stresses in vegetative tissues. Despite the limited role of DME in regulating DNA methylation in vegetative tissues, the dme mutants showed increased susceptibility to bacterial and fungal pathogens. Our study highlights the important functions of DME in vegetative tissues and provides valuable genetic tools for future investigations of DNA demethylation in plants.
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    Histone deacetylase HDA710 controls salt tolerance by regulating ABA signaling in rice
    Farhan Ullah, Qiutao Xu, Yu Zhao and Dao‐Xiu Zhou
    J Integr Plant Biol 2021, 63 (3): 451-467.  
    doi: 10.1111/jipb.13042
    Abstract (Browse 677)  |   Save
    Plants have evolved numerous mechanisms that assist them in withstanding environmental stresses. Histone deacetylases (HDACs) play crucial roles in plant stress responses; however, their regulatory mechanisms remain poorly understood. Here, we explored the function of HDA710/OsHDAC2, a member of the HDAC RPD3/HDA1 family, in stress tolerance in rice (Oryza sativa). We established that HDA710 localizes to both the nucleus and cytoplasm and is involved in regulating the acetylation of histone H3 and H4, specifically targeting H4K5 and H4K16 under normal conditions. HDA710 transcript accumulation levels were strongly induced by abiotic stresses including drought and salinity, as well as by the phytohormones jasmonic acid (JA) and abscisic acid (ABA). hda710 knockout mutant plants showed enhanced salinity tolerance and reduced ABA sensitivity, whereas transgenic plants overexpressing HDA710 displayed the opposite phenotypes. Moreover, ABA‐ and salt‐stress‐responsive genes, such as OsLEA3, OsABI5, OsbZIP72, and OsNHX1, were upregulated in hda710 compared with wild‐type plants. These expression differences corresponded with higher levels of histone H4 acetylation in gene promoter regions in hda710 compared with the wild type under ABA and salt‐stress treatment. Collectively, these results suggest that HDA710 is involved in regulating ABA‐ and salt‐stress‐responsive genes by altering H4 acetylation levels in their promoters.
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    The calcium‐dependent protein kinase ZmCDPK7 functions in heat‐stress tolerance in maize
    Yulong Zhao, Hanwei Du, Yankai Wang, Huali Wang, Shaoyu Yang, Chaohai Li, Ning Chen, Hao Yang, Yihao Zhang, Yulin Zhu, Luyao Yang and Xiuli Hu
    J Integr Plant Biol 2021, 63 (3): 510-527.  
    doi: 10.1111/jipb.13056
    Abstract (Browse 544)  |   Save
    Global warming poses a serious threat to crops. Calcium‐dependent protein kinases (CDPKs)/CPKs play vital roles in plant stress responses, but their exact roles in plant thermotolerance remains elusive. Here, we explored the roles of heat‐induced ZmCDPK7 in thermotolerance in maize. ZmCDPK7‐overexpressing maize plants displayed higher thermotolerance, photosynthetic rates, and antioxidant enzyme activity but lower H2O2 and malondialdehyde (MDA) contents than wild‐type plants under heat stress. ZmCDPK7‐knockdown plants displayed the opposite patterns. ZmCDPK7 is attached to the plasma membrane but can translocate to the cytosol under heat stress. ZmCDPK7 interacts with the small heat shock protein sHSP17.4, phosphorylates sHSP17.4 at Ser‐44 and the respiratory burst oxidase homolog RBOHB at Ser‐99, and upregulates their expression. Site‐directed mutagenesis of sHSP17.4 to generate a Ser‐44‐Ala substitution reduced ZmCDPK7's enhancement of catalase activity but enhanced ZmCDPK7's suppression of MDA accumulation in heat‐stressed maize protoplasts. sHSP17.4, ZmCDPK7, and RBOHB were less strongly upregulated in response to heat stress in the abscisic acid‐deficient mutant vp5 versus the wild type. Pretreatment with an RBOH inhibitor suppressed sHSP17.4 and ZmCDPK7 expression. Therefore, abscisic acid‐induced ZmCDPK7 functions both upstream and downstream of RBOH and participates in thermotolerance in maize by mediating the phosphorylation of sHSP17.4, which might be essential for its chaperone function.
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    Arabidopsis U‐box E3 ubiquitin ligase PUB11 negatively regulates drought tolerance by degrading the receptor‐like protein kinases LRR1 and KIN7
    Xuexue Chen, Tingting Wang, Amin Ur Rehman, Yu Wang, Junsheng Qi, Zhen Li, Chunpeng Song, Baoshan Wang, Shuhua Yang and Zhizhong Gong
    J Integr Plant Biol 2021, 63 (3): 494-509.  
    doi: 10.1111/jipb.13058
    Abstract (Browse 718)  |   Save
    Both plant receptor‐like protein kinases (RLKs) and ubiquitin‐mediated proteolysis play crucial roles in plant responses to drought stress. However, the mechanism by which E3 ubiquitin ligases modulate RLKs is poorly understood. In this study, we showed that Arabidopsis PLANT U‐BOX PROTEIN 11 (PUB11), an E3 ubiquitin ligase, negatively regulates abscisic acid (ABA)‐mediated drought responses. PUB11 interacts with and ubiquitinates two receptor‐like protein kinases, LEUCINE RICH REPEAT PROTEIN 1 (LRR1) and KINASE 7 (KIN7), and mediates their degradation during plant responses to drought stress in vitro and in vivo. pub11 mutants were more tolerant, whereas lrr1 and kin7 mutants were more sensitive, to drought stress than the wild type. Genetic analyses show that the pub11 lrr1 kin7 triple mutant exhibited similar drought sensitivity as the lrr1 kin7 double mutant, placing PUB11 upstream of the two RLKs. Abscisic acid and drought treatment promoted the accumulation of PUB11, which likely accelerates LRR1 and KIN7 degradation. Together, our results reveal that PUB11 negatively regulates plant responses to drought stress by destabilizing the LRR1 and KIN7 RLKs.
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    Protein kinases in plant responses to drought, salt, and cold stress
    Xuexue Chen, Yanglin Ding, Yongqing Yang, Chunpeng Song, Baoshan Wang, Shuhua Yang, Yan Guo and Zhizhong Gong
    J Integr Plant Biol 2021, 63 (1): 53-78.  
    doi: 10.1111/jipb.13061
    Abstract (Browse 1183)  |   Save
    Protein kinases are major players in various signal transduction pathways. Understanding the molecular mechanisms behind plant responses to biotic and abiotic stresses has become critical for developing and breeding climate‐resilient crops. In this review, we summarize recent progress on understanding plant drought, salt, and cold stress responses, with a focus on signal perception and transduction by different protein kinases, especially sucrose nonfermenting1 (SNF1)‐related protein kinases (SnRKs), mitogen‐activated protein kinase (MAPK) cascades, calcium‐dependent protein kinases (CDPKs/CPKs), and receptor‐like kinases (RLKs). We also discuss future challenges in these research fields.
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    Chromatin remodeling factors regulate environmental stress responses in plants
    Ze‐Ting Song, Jian‐Xiang Liu and Jia‐Jia Han
    J Integr Plant Biol 2021, 63 (3): 438-450.  
    doi: 10.1111/jipb.13064
    Abstract (Browse 421)  |   Save
    Environmental stress from climate change and agricultural activity threatens global plant biodiversity as well as crop yield and quality. As sessile organisms, plants must maintain the integrity of their genomes and adjust gene expression to adapt to various environmental changes. In eukaryotes, nucleosomes are the basic unit of chromatin around which genomic DNA is packaged by condensation. To enable dynamic access to packaged DNA, eukaryotes have evolved Snf2 (sucrose nonfermenting 2) family proteins as chromatin remodeling factors (CHRs) that modulate the position of nucleosomes on chromatin. During plant stress responses, CHRs are recruited to specific genomic loci, where they regulate the distribution or composition of nucleosomes, which in turn alters the accessibility of these loci to general transcription or DNA damage repair machinery. Moreover, CHRs interplay with other epigenetic mechanisms, including DNA methylation, histone modifications, and deposition of histone variants. CHRs are also involved in RNA processing at the post‐transcriptional level. In this review, we discuss major advances in our understanding of the mechanisms by which CHRs function during plants’ response to environmental stress.
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    An update on the function and regulation of methylerythritol phosphate and mevalonate pathways and their evolutionary dynamics
    Xiaojun Pu, Xiumei Dong, Qing Li, Zexi Chen and Li Liu
    J Integr Plant Biol 2021, 63 (7): 1211-1226.  
    DOI: 10.1111/jipb.13076
    Abstract (Browse 325)  |   Save
    Isoprenoids are among the largest and most chemically diverse classes of organic compounds in nature and are involved in the processes of photosynthesis, respiration, growth, development, and plant responses to stress. The basic building block units for isoprenoid synthesis—isopentenyl diphosphate and its isomer dimethylallyl diphosphate—are generated by the mevalonate (MVA) and methylerythritol phosphate (MEP) pathways. Here, we summarize recent advances on the roles of the MEP and MVA pathways in plant growth, development and stress responses, and attempt to define the underlying gene networks that orchestrate the MEP and MVA pathways in response to developmental or environmental cues. Through phylogenomic analysis, we also provide a new perspective on the evolution of the plant isoprenoid pathway. We conclude that the presence of the MVA pathway in plants may be associated with the transition from aquatic to subaerial and terrestrial environments, as lineages for its core components are absent in green algae. The emergence of the MVA pathway has acted as a key evolutionary event in plants that facilitated land colonization and subsequent embryo development, as well as adaptation to new and varied environments.
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    SIZ1 negatively regulates aluminum resistance by mediating the STOP1–ALMT1 pathway in Arabidopsis
    Jiameng Xu, Jiayong Zhu, Jiajia Liu, Junxia Wang, Zhaojun Ding and Huiyu Tian
    J Integr Plant Biol 2021, 63 (6): 1147-1160.  
    DOI: 10.1111/jipb.13091
    Abstract (Browse 366)  |   Save
    Sensitive to proton rhizotoxicity 1 (STOP1) functions as a crucial regulator of root growth during aluminum (Al) stress. However, how this transcription factor is regulated by Al stress to affect downstream genes expression is not well understood. To explore the underlying mechanisms of the function and regulation of STOP1, we employed a yeast two hybrid screen to identify STOP1-interacting proteins. The SUMO E3 ligase SIZ1, was found to interact with STOP1 and mainly facilitate its SUMO modification at K40 and K212 residues. Simultaneous introduction of K40R and K212R substitutions in STOP1 enhances its transactivation activity to upregulate the expression of aluminum-activated malate transporter 1 (ALMT1) via increasing the association with mediator 16 (MED16) transcriptional co-activator. Loss of function of SIZ1 causes highly increased expression of ALMT1, thus enhancing Al-induced malate exudation and Al tolerance. Also, we found that the protein level of SIZ1 is reduced in response to Al stress. Genetic evidence demonstrates that STOP1/ALMT1 is epistatic to SIZ1 in regulating root growth response to Al stress. This study suggests a mechanism about how the SIZ1–STOP1–ALMT1 signaling module is involved in root growth response to Al stress.
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    Drought stress and plant ecotype drive microbiome recruitment in switchgrass rhizosheath
    Tie‐Yuan Liu, Nenghui Ye, Xinyu Wang, Debatosh Das, Yuxiang Tan, Xiangkai You, Mingxiu Long, Tianming Hu, Lei Dai, Jianhua Zhang and Mo‐Xian Chen
    J Integr Plant Biol 2021, 63 (10): 1753-1774.  
    doi: 10.1111/jipb.13154
    Abstract (Browse 295)  |   Save
    The rhizosheath, a layer of soil grains that adheres firmly to roots, is beneficial for plant growth and adaptation to drought environments. Switchgrass is a perennial C4 grass which can form contact rhizosheath under drought conditions. In this study, we characterized the microbiomes of four different rhizocompartments of two switchgrass ecotypes (Alamo and Kanlow) grown under drought or well-watered conditions via 16S ribosomal RNA amplicon sequencing. These four rhizocompartments, the bulk soil, rhizosheath soil, rhizoplane, and root endosphere, harbored both distinct and overlapping microbial communities. The root compartments (rhizoplane and root endosphere) displayed low-complexity communities dominated by Proteobacteria and Firmicutes. Compared to bulk soil, Cyanobacteria and Bacteroidetes were selectively enriched, while Proteobacteria and Firmicutes were selectively depleted, in rhizosheath soil. Taxa from Proteobacteria or Firmicutes were specifically selected in Alamo or Kanlow rhizosheath soil. Following drought stress, Citrobacter and Acinetobacter were further enriched in rhizosheath soil, suggesting that rhizosheath microbiome assembly is driven by drought stress. Additionally, the ecotype-specific recruitment of rhizosheath microbiome reveals their differences in drought stress responses. Collectively, these results shed light on rhizosheath microbiome recruitment in switchgrass and lay the foundation for the improvement of drought tolerance in switchgrass by regulating the rhizosheath microbiome.
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    The direct targets of CBFs: In cold stress response and beyond
    Yue Song, Xiaoyan Zhang, Minze Li, Hao Yang, Diyi Fu, Jian Lv, Yanglin Ding, Zhizhong Gong, Yiting Shi and Shuhua Yang
    J Integr Plant Biol 2021, 63 (11): 1874-1887.  
    DOI: 10.1111/jipb.13161
    Abstract (Browse 533)  |   Save
    Cold acclimation in Arabidopsis thaliana triggers a significant transcriptional reprogramming altering the expression patterns of thousands of cold-responsive (COR) genes. Essential to this process is the C-repeat binding factor (CBF)-dependent pathway, involving the activity of AP2/ERF (APETALA2/ethylene-responsive factor)-type CBF transcription factors required for plant cold acclimation. In this study, we performed chromatin immunoprecipitation assays followed by deep sequencing (ChIP-seq) to determine the genome-wide binding sites of the CBF transcription factors. Cold-induced CBF proteins specifically bind to the conserved C-repeat (CRT)/dehydration-responsive elements (CRT/DRE; G/ACCGAC) of their target genes. A Gene Ontology enrichment analysis showed that 1,012 genes are targeted by all three CBFs. Combined with a transcriptional analysis of the cbf1,2,3 triple mutant, we define 146 CBF regulons as direct CBF targets. In addition, the CBF-target genes are significantly enriched in functions associated with hormone, light, and circadian rhythm signaling, suggesting that the CBFs act as key integrators of endogenous and external environmental cues. Our findings not only define the genome-wide binding patterns of the CBFs during the early cold response, but also provide insights into the role of the CBFs in regulating multiple biological processes of plants.
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