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OsEDM2L mediates m6A of EAT1 transcript for proper alternative splicing and polyadenylation regulating rice tapetal degradation
Kun Ma, Jingluan Han, Zixu Zhang, Heying Li, Yanchang Zhao, Qinlong Zhu, Yongyao Xie, Yao‐Guang Liu and Letian Chen
J Integr Plant Biol 2021, 63 (11): 1982-1994.  
DOI: 10.1111/jipb.13167
Abstract (Browse 1426)  |   Save
N6-methyladenosine (m6A) modification affects the post-transcriptional regulation of eukaryotic gene expression, but the underlying mechanisms and their effects in plants remain largely unknown. Here, we report that the N6-adenine methyltransferase-like domain-containing protein ENHANCED DOWNY MILDEW 2-LIKE (OsEDM2L) is essential for rice (Oryza sativa L.) anther development. The osedm2l knockout mutant showed delayed tapetal programmed cell death (PCD) and defective pollen development. OsEDM2L interacts with the transcription factors basic helix-loop-helix 142 and TAPETUM DEGENERATION RETARDATION to regulate the expression of ETERNAL TAPETUM 1 (EAT1), a positive regulator of tapetal PCD. Mutation of OsEDM2L altered the transcriptomic m6A landscape, and caused a distinct m6A modification of the EAT1 transcript leading to dysregulation of its alternative splicing and polyadenylation, followed by suppression of the EAT1 target genes OsAP25 and OsAP37 for tapetal PCD. Therefore, OsEDM2L is indispensable for proper messenger RNA m6A modification in rice anther development.
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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 1062)  |   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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Cited: Web of Science(123)
  
Single-cell RNA sequencing reveals a high-resolution cell atlas of xylem in Populus
Hui Li, Xinren Dai, Xiong Huang, Mengxuan Xu, Qiao Wang, Xiaojing Yan, Ronald R. Sederoff and Quanzi Li
J Integr Plant Biol 2021, 63 (11): 1906-1921.  
DOI: 10.1111/jipb.13159
Abstract (Browse 1001)  |   Save
High-throughput single-cell RNA sequencing (scRNA-seq) has advantages over traditional RNA-seq to explore spatiotemporal information on gene dynamic expressions in heterogenous tissues. We performed Drop-seq, a method for the dropwise sequestration of single cells for sequencing, on protoplasts from the differentiating xylem of Populus alba×Populus glandulosa. The scRNA-seq profiled 9,798 cells, which were grouped into 12 clusters. Through characterization of differentially expressed genes in each cluster and RNA in situ hybridizations, we identified vessel cells, fiber cells, ray parenchyma cells and xylem precursor cells. Diffusion pseudotime analyses revealed the differentiating trajectory of vessels, fiber cells and ray parenchyma cells and indicated a different differentiation process between vessels and fiber cells, and a similar differentiation process between fiber cells and ray parenchyma cells. We identified marker genes for each cell type (cluster) and key candidate regulators during developmental stages of xylem cell differentiation. Our study generates a high-resolution expression atlas of wood formation at the single cell level and provides valuable information on wood formation.
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The ScCas9++ variant expands the CRISPR toolbox for genome editing in plants
Taoli Liu, Dongchang Zeng, Zhiye Zheng, Zhansheng Lin, Yang Xue, Tie Li, Xianrong Xie, Genglu Ma, Yao‐Guang Liu and Qinlong Zhu
J Integr Plant Biol 2021, 63 (9): 1611-1619.  
doi: 10.1111/jipb.13164
Abstract (Browse 953)  |   Save
The development of clustered regularly interspaced palindromic repeats (CRISPR)-associated protein (Cas) variants with a broader recognition scope is critical for further improvement of CRISPR/Cas systems. The original Cas9 protein from Streptococcus canis (ScCas9) can recognize simple NNG-protospacer adjacent motif (PAM) targets, and therefore possesses a broader range relative to current CRISPR/Cas systems, but its editing efficiency is low in plants. Evolved ScCas9+ and ScCas9++ variants have been shown to possess higher editing efficiencies in human cells, but their activities in plants are currently unknown. Here, we utilized codon-optimized ScCas9, ScCas9+ and ScCas9++ and a nickase variant ScCas9n++ to systematically investigate genome cleavage activity and cytidine base editing efficiency in rice (Oryza sativa L.). This analysis revealed that ScCas9++ has higher editing efficiency than ScCas9 and ScCas9+ in rice. Furthermore, we fused the evolved cytidine deaminase PmCDA1 with ScCas9n++ to generate a new evoBE4max-type cytidine base editor, termed PevoCDA1-ScCas9n++. This base editor achieved stable and efficient multiplex-site base editing at NNG-PAM sites with wider editing windows (C1–C17) and without target sequence context preference. Multiplex-site base editing of the rice genes OsWx (three targets) and OsEui1 (two targets) achieved simultaneous editing and produced new rice germplasm. Taken together, these results demonstrate that ScCas9++ represents a crucial new tool for improving plant editing.
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Plant abiotic stress: New insights into the factors that activate and modulate plant responses
Zhizhong Gong
J Integr Plant Biol 2021, 63 (3): 429-430.  
doi: 10.1111/jipb.13079
Abstract (Browse 775)  |   Save

Plants experience a remarkable range of interactions with the abiotic factors in their environments, both aboveground (light, temperature, mechanical stress) and belowground (soil moisture, nutrients, and mechanical properties). Plants’ abilities to sense diverse environmental parameters and initiate signaling pathways that activate precise responses have crucial implications for their survival. Plant abiotic interactions also remain a fascinating area of research, providing basic insight into plant biology and enabling efforts to improve crop plants.

Transcriptional and epigenetic regulation mediate many abiotic stress responses. For example, Ullah et al. (2021) reveal a role for histone deacetylases in modulating genes that respond to abscisic acid (ABA) and salt stress. Moreover, Liu et al. (2021) compared the pools of nascent RNAs and messenger RNAs during heat shock in Arabidopsis and identified effects that occurred at different steps in transcription, from recruitment of polymerase to termination. In this issue, Song et al. (2021) review the functions of chromatin‐remodeling factors in plant stress responses and Chang et al. (2020) provide a comprehensive overview of epigenetic regulation in plant abiotic stress responses.

Kinases have key functions in abiotic stress signaling (reviewed in Chen et al., 2021a) and emerging research shows that plants modulate kinase activities to rein in stress responses. For example, Yu et al. (2021a) explore the function of the Arabidopsis mitogen‐activated protein kinase kinase kinase MAPKKK18 in drought stress, finding that two RING finger ubiquitin ligases control MAPKKK18 protein levels, thus negatively regulating its role in drought stress responses. Moreover, Chen et al. (2021b) identify two Arabidopsis PLANT U‐BOX (PUB) ubiquitin ligases that also negatively modulate drought responses by ubiquitin‐mediated degradation of the receptor‐like protein kinases LEUCINE‐RICH REPEAT PROTEIN1 and KINASE7. Exploring the role of calcium‐dependent protein kinases, Zhao et al. (2021) find that maize (Zea mays) ZmCDPK7 translocates from the plasma membrane to the cytosol under heat stress, where it interacts with a heat shock protein and a Respiratory Burst Oxidase Homolog protein to regulate reactive oxygen species (ROS) in thermotolerance. Understanding how plant cells regulate kinase activities to optimize their stress responses will improve our ability to enhance plant stress tolerance.

In addition to kinases, other post‐translational factors are emerging as important, rapid mechanisms to regulate abiotic stress responses. In addition to modulating kinase levels, the ubiquitin‐mediated proteolysis system functions in drought stress tolerance: Yu et al. (2021b) identify a RING‐type E3 Ub ligase that localizes to the endoplasmic reticulum and participates in mitigating drought stress‐induced proteotoxic stress. Intriguingly, Chu et al. (2021) show that an Arabidopsis Type 2C protein phosphatase regulates the activity of a high‐affinity K+‐transporter to modulate sodium levels under salt stress conditions.

New findings are expanding our understanding of the mechanisms by which the “drought stress” hormone ABA acts (reviewed in Chen et al., 2020). For example, Ma et al. (2021) discovered that the ABA receptors increase the activity of the ABA‐conjugating enzyme UDP‐glucosyltransferase in a rapid mechanism to alter ABA levels. Qiao et al. (2021) explored the crosstalk between ABA and ethylene in ripening of common fig (Ficus carica). Ngoc et al. (2021) show that N4‐methylcytidine (m4C) modification of the 16S chloroplast ribosomal RNA participates in the Arabidopsis response to ABA. The identification of these diverse cellular mechanisms affecting ABA signaling give us more information on this key hormone.

Root architecture and root growth affect plant tolerance to multiple stresses, particularly drought and nutrient deficiency. An intriguing study by Wang et al. (2021) examines the function of salicylic acid (SA), a key hormone for plant immunity, in maintenance of the root quiescent center (QC), finding that SA promotes ROS production and cell division in the QC. Moreover, Su et al. (2021) reveal that SUCROSE NON‐FERMENTING kinases 2 function downstream of ABA to modulate the activity of the nitrate transceptor NITRATE TRANSPORTER1.1 and thus root growth under nitrogen deficiency, suggesting that ABA signaling has a cross‐talk with nitrate uptaking in plants.

Understanding the mechanisms that mediate abiotic stress tolerance has key implications for improving crops, as the sustainable crops of the future will require tolerance to multiple stresses, including more than one stress at a time. Translating these exciting results from controlled laboratory conditions to the field remains a major challenge. Moreover, approaches such as removing a negative regulator, or activating positive regulators, will likely have knock‐on effects on plant fitness and yield. The papers in this issue provide important segments of the broad, foundational knowledge that will be required to design the stress‐tolerant, high‐yield crops of the future.

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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 737)  |   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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Genome editing for plant research and crop improvement
Xiangqiang Zhan, Yuming Lu, Jian-Kang Zhu and Jose Ramon Botella
J Integr Plant Biol 2021, 63 (1): 3-33.  
doi: 10.1111/jipb.13063
Abstract (Browse 693)  |   Save
The advent of clustered regularly interspaced short palindromic repeat (CRISPR) has had a profound impact on plant biology, and crop improvement. In this review, we summarize the state‐of‐the‐art development of CRISPR technologies and their applications in plants, from the initial introduction of random small indel (insertion or deletion) mutations at target genomic loci to precision editing such as base editing, prime editing and gene targeting. We describe advances in the use of class 2, types II, V, and VI systems for gene disruption as well as for precise sequence alterations, gene transcription, and epigenome control.
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Cited: Web of Science(31)
  
Contribution of phenylpropanoid metabolism to plant development and plant–environment interactions
Nai-Qian Dong and Hong-Xuan Lin
J Integr Plant Biol 2021, 63 (1): 180-209.  
doi: 10.1111/jipb.13054
Abstract (Browse 642)  |   Save
Phenylpropanoid metabolism is one of the most important metabolisms in plants, yielding more than 8,000 metabolites contributing to plant development and plant–environment interplay. Phenylpropanoid metabolism materialized during the evolution of early freshwater algae that were initiating terrestrialization and land plants have evolved multiple branches of this pathway, which give rise to metabolites including lignin, flavonoids, lignans, phenylpropanoid esters, hydroxycinnamic acid amides, and sporopollenin. Recent studies have revealed that many factors participate in the regulation of phenylpropanoid metabolism, and modulate phenylpropanoid homeostasis when plants undergo successive developmental processes and are subjected to stressful environments. In this review, we summarize recent progress on elucidating the contribution of phenylpropanoid metabolism to the coordination of plant development and plant–environment interaction, and metabolic flux redirection among diverse metabolic routes. In addition, our review focuses on the regulation of phenylpropanoid metabolism at the transcriptional, post‐transcriptional, post‐translational, and epigenetic levels, and in response to phytohormones and biotic and abiotic stresses.
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Cited: Web of Science(186)
  
Arabidopsis E3 ligase KEG associates with and ubiquitinates MKK4 and MKK5 to regulate plant immunity
Chenyang Gao, Pengwei Sun, Wei Wang and Dingzhong Tang
J Integr Plant Biol 2021, 63 (2): 327-339.  
doi: 10.1111/jipb.13007
Abstract (Browse 611)  |   Save
Mitogen‐activated protein kinase (MAPK) cascades are highly conserved signaling modules that regulate plant immune responses. The Arabidopsis thaliana Raf‐like MAPK kinase kinase ENHANCED DISEASE RESISTANCE1 (EDR1) is a key negative regulator of plant immunity that affects the protein levels of MKK4 and MKK5, two important MAPK cascade members, but the underlying mechanism is poorly understood. Here, genome‐wide phosphorylation analysis demonstrated that the E3 ligase KEEP ON GOING (KEG) is phosphorylated in the edr1 mutant but not the wild type, suggesting that EDR1 negatively affects KEG phosphorylation. The identified phosphorylation sites in KEG appear to be important for its accumulation. The keg‐4 mutant, a previously identified edr1 suppressor, enhances susceptibility to the powdery mildew pathogen Golovinomyces cichoracearum. In addition, MKK4 and MKK5 protein levels are reduced in the keg‐4 mutant. Furthermore, we demonstrate that MKK4 and MKK5 associate with full‐length KEG, but not with truncated KEG‐RK or KEG‐RKA, and that KEG ubiquitinates and mediates the degradation of MKK4 and MKK5. Taken together, these results indicate that MKK4 and MKK5 protein levels are regulated by KEG via ubiquitination, uncovering a mechanism by which plants fine‐tune immune responses by regulating the homeostasis of key MAPK cascade members via ubiquitination and degradation.
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Cited: Web of Science(29)
  
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 569)  |   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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Phyllosphere microbiota: Community dynamics and its interaction with plant hosts
Tianyu Gong and Xiu‐Fang Xin
J Integr Plant Biol 2021, 63 (2): 297-304.  
doi: 10.1111/jipb.13060
Abstract (Browse 561)  |   Save
Plants are colonized by various microorganisms in natural environments. While many studies have demonstrated key roles of the rhizosphere microbiota in regulating biological processes such as nutrient acquisition and resistance against abiotic and biotic challenges, less is known about the role of the phyllosphere microbiota and how it is established and maintained. This review provides an update on current understanding of phyllosphere community assembly and the mechanisms by which plants and microbes establish the phyllosphere microbiota for plant health.
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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 550)  |   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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A global alfalfa diversity panel reveals genomic selection signatures in Chinese varieties and genomic associations with root development
Lin Chen, Fei He, Ruicai Long, Fan Zhang, Mingna Li, Zhen Wang, Junmei Kang and Qingchuan Yang
J Integr Plant Biol 2021, 63 (11): 1937-1951.  
DOI: 10.1111/jipb.13172
Abstract (Browse 538)  |   Save
Alfalfa (Medicago sativa L.) is an important forage crop worldwide. However, little is known about the effects of breeding status and different geographical populations on alfalfa improvement. Here, we sequenced 220 alfalfa core germplasms and determined that Chinese alfalfa cultivars form an independent group, as evidenced by comparisons of FST values between different subgroups, suggesting that geographical origin plays an important role in group differentiation. By tracing the influence of geographical regions on the genetic diversity of alfalfa varieties in China, we identified 350 common candidate genetic regions and 548 genes under selection. We also defined 165 loci associated with 24 important traits from genome-wide association studies. Of those, 17 genomic regions closely associated with a given phenotype were under selection, with the underlying haplotypes showing significant differences between subgroups of distinct geographical origins. Based on results from expression analysis and association mapping, we propose that 6-phosphogluconolactonase (MsPGL) and a gene encoding a protein with NHL domains (MsNHL) are critical candidate genes for root growth. In conclusion, our results provide valuable information for alfalfa improvement via molecular breeding.
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Reciprocal regulation between the negative regulator PP2CG1 phosphatase and the positive regulator OST1 kinase confers cold response in Arabidopsis
Jian Lv, Jingyan Liu, Yuhang Ming, Yiting Shi, Chunpeng Song, Zhizhong Gong, Shuhua Yang and Yanglin Ding
J Integr Plant Biol 2021, 63 (8): 1568-1587.  
DOI: 10.1111/jipb.13100
Abstract (Browse 531)  |   Save
Protein phosphorylation and dephosphorylation have been reported to play important roles in plant cold responses. In addition, phospho-regulatory feedback is a conserved mechanism for biological processes and stress responses in animals and plants. However, it is less well known that a regulatory feedback loop is formed by the protein kinase and the protein phosphatase in plant responses to cold stress. Here, we report that OPEN STOMATA 1 (OST1) and PROTEIN PHOSPHATASE 2C G GROUP 1 (PP2CG1) reciprocally regulate the activity during the cold stress response. The interaction of PP2CG1 and OST1 is inhibited by cold stress, which results in the release of OST1 at the cytoplasm and nucleus from suppression by PP2CG1. Interestingly, cold-activated OST1 phosphorylates PP2CG1 to suppress its phosphatase activity, thereby amplifying cold signaling in plants. Mutations of PP2CG1 and its homolog PP2CG2 enhance freezing tolerance, whereas overexpression of PP2CG1 decreases freezing tolerance. Moreover, PP2CG1 negatively regulates protein levels of C-REPEAT BINDING FACTORs (CBFs) under cold stress. Our results uncover a phosphor/dephosphor-regulatory feedback loop mediated by PP2CG1 phosphatase and OST1 protein kinase in plant cold responses.
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Co-regulation of indole glucosinolates and camalexin biosynthesis by CPK5/CPK6 and MPK3/MPK6 signaling pathways
Liuyi Yang, Yan Zhang, Rongxia Guan, Sen Li, Xuwen Xu, Shuqun Zhang and Juan Xu
J Integr Plant Biol 2020, 62 (11): 1780-1796.  
doi: 10.1111/jipb.12973
Abstract (Browse 523)  |   Save

Secondary plant metabolites, represented by indole glucosinolates (IGS) and camalexin, play important roles in Arabidopsis immunity. Previously, we demonstrated the importance of MPK3 and MPK6, two closely related MAPKs, in regulating Botrytis cinerea (Bc)‐induced IGS and camalexin biosynthesis. Here we report that CPK5 and CPK6, two redundant calcium‐dependent protein kinases (CPKs), are also involved in regulating the biosynthesis of these secondary metabolites. The loss‐of‐function of both CPK5 and CPK6 compromises plant resistance to Bc. Expression profiling of CPK5‐VK transgenic plants, in which a truncated constitutively active CPK5 is driven by a steroid‐inducible promoter, revealed that biosynthetic genes of both IGS and camalexin pathways are coordinately upregulated after the induction of CPK5‐VK, leading to high‐level accumulation of camalexin and 4‐methoxyindole‐3‐yl‐methylglucosinolate (4MI3G). Induction of camalexin and 4MI3G, as well as the genes in their biosynthesis pathways, is greatly compromised in cpk5 cpk6 mutant in response to Bc. In a conditional cpk5 cpk6 mpk3 mpk6 quadruple mutant, Bc resistance and induction of IGS and camalexin are further reduced in comparison to either cpk5 cpk6 or conditional mpk3 mpk6 double mutant, suggesting that both CPK5/CPK6 and MPK3/MPK6 signaling pathways contribute to promote the biosynthesis of 4MI3G and camalexin in defense against Bc.

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Abscisic acid signaling negatively regulates nitrate uptake via phosphorylation of NRT1.1 by SnRK2s in Arabidopsis
Hang Su, Tian Wang, Chuanfeng Ju, Jinping Deng, Tianqi Zhang, Mengjiao Li, Hui Tian and Cun Wang
J Integr Plant Biol 2021, 63 (3): 597-610.  
doi: 10.1111/jipb.13057
Abstract (Browse 501)  |   Save
Nitrogen (N) is a limiting nutrient for plant growth and productivity. The phytohormone abscisic acid (ABA) has been suggested to play a vital role in nitrate uptake in fluctuating N environments. However, the molecular mechanisms underlying the involvement of ABA in N deficiency responses are largely unknown. In this study, we demonstrated that ABA signaling components, particularly the three subclass III SUCROSE NON‐FERMENTING1 (SNF1)‐RELATED PROTEIN KINASE 2S (SnRK2) proteins, function in root foraging and uptake of nitrate under N deficiency in Arabidopsis thaliana. The snrk2.2snrk2.3snrk2.6 triple mutant grew a longer primary root and had a higher rate of nitrate influx and accumulation compared with wild‐type plants under nitrate deficiency. Strikingly, SnRK2.2/2.3/2.6 proteins interacted with and phosphorylated the nitrate transceptor NITRATE TRANSPORTER1.1 (NRT1.1) in vitro and in vivo. The phosphorylation of NRT1.1 by SnRK2s resulted in a significant decrease of nitrate uptake and impairment of root growth. Moreover, we identified NRT1.1Ser585 as a previously unknown functional site: the phosphomimetic NRT1.1S585D was impaired in both low‐ and high‐affinity transport activities. Taken together, our findings provide new insight into how plants fine‐tune growth via ABA signaling under N deficiency.
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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 489)  |   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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PP2A interacts with KATANIN to promote microtubule organization and conical cell morphogenesis
Huibo Ren, Jinqiu Rao, Min Tang, Yaxing Li, Xie Dang and Deshu Lin
J Integr Plant Biol 2022, 64 (8): 1514-1530.  
DOI: 10.1111/jipb.13281
Abstract (Browse 488)  |   Save

The organization of the microtubule cytoskeleton is critical for cell and organ morphogenesis. The evolutionarily conserved microtubule-severing enzyme KATANIN plays critical roles in microtubule organization in the plant and animal kingdoms. We previously used conical cell of Arabidopsis thaliana petals as a model system to investigate cortical microtubule organization and cell morphogenesis and determined that KATANIN promotes the formation of circumferential cortical microtubule arrays in conical cells. Here, we demonstrate that the conserved protein phosphatase PP2A interacts with and dephosphorylates KATANIN to promote the formation of circumferential cortical microtubule arrays in conical cells. KATANIN undergoes cycles of phosphorylation and dephosphorylation. Using co-immunoprecipitation coupled with mass spectrometry, we identified PP2A subunits as KATANIN-interacting proteins. Further biochemical studies showed that PP2A interacts with and dephosphorylates KATANIN to stabilize its cellular abundance. Similar to the katanin mutant, mutants for genes encoding PP2A subunits showed disordered cortical microtubule arrays and defective conical cell shape. Taken together, these findings identify PP2A as a regulator of conical cell shape and suggest that PP2A mediates KATANIN phospho-regulation during plant cell morphogenesis.

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Allele-aware chromosome-scale assembly of the allopolyploid genome of hexaploid Ma bamboo (Dendrocalamus latiflorus Munro)
Yushan Zheng, Deming Yang, Jundong Rong, Liguang Chen, Qiang Zhu, Tianyou He, Lingyan Chen, Jing Ye, Lili Fan, Yubang Gao, Hangxiao Zhang and Lianfeng Gu
J Integr Plant Biol 2022, 64 (3): 649-670.  
doi: 10.1111/jipb.13217
Abstract (Browse 488)  |   Save
Dendrocalamus latiflorus Munro is a woody clumping bamboo with rapid shoot growth. Both genetic transformation and clustered regularly interspaced short palindromic repeats (CRISPR)/CRISPR-associated protein 9 (Cas9) gene editing techniques are available for D. latiflorus, enabling reverse genetic approaches. Thus, D. latiflorus has the potential to be a model bamboo species. However, the genome sequence of D. latiflorus has remained unreported due to its polyploidy and large genome size. Here, we sequenced the D. latiflorus genome and assembled it into three allele-aware subgenomes (AABBCC), representing the largest genome of a major bamboo species. We assembled 70 allelic chromosomes (2, 737 Mb) for hexaploid D. latiflorus using both single-molecule sequencing from the Pacific Biosciences (PacBio) Sequel platform and chromosome conformation capture sequencing (Hi-C). Repetitive sequences comprised 52.65% of the D. latiflorus genome. We annotated 135 231 protein-coding genes in the genome based on transcriptomes from eight different tissues. Transcriptome sequencing using RNA-Seq and PacBio single-molecule real-time long-read isoform sequencing revealed highly differential alternative splicing (AS) between non-abortive and abortive shoots, suggesting that AS regulates the abortion rate of bamboo shoots. This high-quality hexaploid genome and comprehensive strand-specific transcriptome datasets for this Poaceae family member will pave the way for bamboo research using D. latiflorus as a model species.
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A histone H3K27me3 reader cooperates with a family of PHD finger‐containing proteins to regulate flowering time in Arabidopsis
Feng Qian, Qiu‐Yuan Zhao, Tie‐Nan Zhang, Yu‐Lu Li, Yin‐Na Su, Lin Li, Jian‐Hua Sui, She Chen and Xin‐Jian He
J Integr Plant Biol 2021, 63 (4): 787-802.  
DOI: 10.1111/jipb.13067
Abstract (Browse 487)  |   Save
Trimethylated histone H3 lysine 27 (H3K27me3) is a repressive histone marker that regulates a variety of developmental processes, including those that determine flowering time. However, relatively little is known about the mechanism of how H3K27me3 is recognized to regulate transcription. Here, we identified BAH domain‐containing transcriptional regulator 1 (BDT1) as an H3K27me3 reader. BDT1 is responsible for preventing flowering by suppressing the expression of flowering genes. Mutation of the H3K27me3 recognition sites in the BAH domain disrupted the binding of BDT1 to H3K27me3, leading to de‐repression of H3K27me3‐enriched flowering genes and an early‐flowering phenotype. We also found that BDT1 interacts with a family of PHD finger‐containing proteins, which we named PHD1–6, and with CPL2, a Pol II carboxyl terminal domain (CTD) phosphatase responsible for transcriptional repression. Pull‐down assays showed that the PHD finger‐containing proteins can enhance the binding of BDT1 to the H3K27me3 peptide. Mutations in all of the PHD genes caused increased expression of flowering genes and an early‐flowering phenotype. This study suggests that the binding of BDT1 to the H3K27me3 peptide, which is enhanced by PHD proteins, is critical for preventing early flowering.
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Potassium and phosphorus transport and signaling in plants
Yi Wang, Yi-Fang Chen and Wei-Hua Wu
J Integr Plant Biol 2021, 63 (1): 34-52.  
doi: 10.1111/jipb.13053
Abstract (Browse 484)  |   Save
Nitrogen (N), potassium (K), and phosphorus (P) are essential macronutrients for plant growth and development, and their availability affects crop yield. Compared with N, the relatively low availability of K and P in soils limits crop production and thus threatens food security and agricultural sustainability. Improvement of plant nutrient utilization efficiency provides a potential route to overcome the effects of K and P deficiencies. Investigation of the molecular mechanisms underlying how plants sense, absorb, transport, and use K and P is an important prerequisite to improve crop nutrient utilization efficiency. In this review, we summarize current understanding of K and P transport and signaling in plants, mainly taking Arabidopsis thaliana and rice (Oryza sativa) as examples. We also discuss the mechanisms coordinating transport of N and K, as well as P and N.
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Cited: Web of Science(72)
  
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 483)  |   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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Cited: Web of Science(116)
  
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 476)  |   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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Cited: Web of Science(29)
  
Targeted mutagenesis in Arabidopsis thaliana using CRISPR-Cas12b/C2c1
Fan Wu , Xinyu Qiao , Yafei Zhao , Ziyi Zhang , Yifan Gao, Lingfeng Shi , Haokun Du , Lulu Wang , Ya‐Jie Zhang , Yu Zhang , Langyu Liu, Quan Wang and Dejing Kong
J Integr Plant Biol 2020, 62 (11): 1653-1658.  
doi: 10.1111/jipb.12944
Abstract (Browse 474)  |   Save

Cas12b/C2c1 is a newly identified class 2 CRISPR endonuclease that was recently engineered for targeted genome editing in mammals and rice. To explore the potential applications of the CRISPR‐Cas12b system in the dicot Arabidopsis thaliana, we selected BvCas12b and BhCas12b v4 for analysis. We successfully used both endonucleases to induce mutations, perform multiplex genome editing, and create large deletions at multiple loci. No significant mutations were detected at potential off‐target sites. Analysis of the insertion/deletion frequencies and patterns of mutants generated via targeted gene mutagenesis highlighted the potential utility of CRISPR‐Cas12b systems for genome editing in Arabidopsis.

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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 470)  |   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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FT5a interferes with the Dt1-AP1 feedback loop to control flowering time and shoot determinacy in soybean
Lin Yue, Xiaoming Li, Chao Fang, Liyu Chen, Hui Yang, Jie Yang, Zhonghui Chen, Haiyang Nan, Linnan Chen, Yuhang Zhang, Haiyang Li, Xingliang Hou, Zhicheng Dong, James L. Weller, Jun Abe, Baohui Liu and Fanjiang Kong
J Integr Plant Biol 2021, 63 (6): 1004-1020.  
doi: 10.1111/jipb.13070
Abstract (Browse 463)  |   Save
Flowering time and stem growth habit determine inflorescence architecture in soybean, which in turn influences seed yield. Dt1, a homolog of Arabidopsis TERMINAL FLOWER 1 (TFL1), is a major controller of stem growth habit, but its underlying molecular mechanisms remain unclear. Here, we demonstrate that Dt1 affects node number and plant height, as well as flowering time, in soybean under long-day conditions. The bZIP transcription factor FDc1 physically interacts with Dt1, and the FDc1-Dt1 complex directly represses the expression of APETALA1 (AP1). We propose that FT5a inhibits Dt1 activity via a competitive interaction with FDc1 and directly upregulates AP1. Moreover, AP1 represses Dt1 expression by directly binding to the Dt1 promoter, suggesting that AP1 and Dt1 form a suppressive regulatory feedback loop to determine the fate of the shoot apical meristem. These findings provide novel insights into the roles of Dt1 and FT5a in controlling the stem growth habit and flowering time in soybean, which determine the adaptability and grain yield of this important crop.
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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 453)  |   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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Receptor‐like kinases MDS1 and MDS2 promote SUMM2‐mediated immunity
Yanan Liu, Xionghui Zhong, Zhibin Zhang, Jiameng Lan, Xingchuan Huang, Hainan Tian, Xin Li and Yuelin Zhang
J Integr Plant Biol 2021, 63 (2): 277-282.  
doi: 10.1111/jipb.12978
Abstract (Browse 452)  |   Save
Disruption of the MEKK1‐MKK1/MKK2‐MPK4 kinase cascade leads to activation of immunity mediated by the nucleotide‐binding leucine‐rich repeat (NLR) immune receptor SUMM2, which monitors the phosphorylation status of CRCK3. Here we report that two receptor‐like kinases (RLKs), MDS1, and MDS2, function redundantly to promote SUMM2‐mediated immunity. Activation of SUMM2‐mediated immunity is dependent on MDS1, and to a less extent on MDS2. MDS1 associates with CRCK3 in planta and can phosphorylate CRCK3 in vitro, suggesting that it may target CRCK3 to positively regulate SUMM2‐mediated signaling. Our finding highlights a new defense mechanism where RLKs promote NLR‐mediated immunity.
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Recent progression and future perspectives in cotton genomic breeding
Zhaoen Yang, Chenxu Gao, Yihao Zhang, Qingdi Yan, Wei Hu, Lan Yang, Zhi Wang and Fuguang Li
J Integr Plant Biol 2023, 65 (2): 548-569.  
doi: 10.1111/jipb.13388
Abstract (Browse 444)  |   Save
Upland cotton is an important global cash crop for its long seed fibers and high edible oil and protein content. Progress in cotton genomics promotes the advancement of cotton genetics, evolutionary studies, functional genetics, and breeding, and has ushered cotton research and breeding into a new era. Here, we summarize high-impact genomics studies for cotton from the last 10 years. The diploid Gossypium arboreum and allotetraploid Gossypium hirsutum are the main focus of most genetic and genomic studies. We next review recent progress in cotton molecular biology and genetics, which builds on cotton genome sequencing efforts, population studies, and functional genomics, to provide insights into the mechanisms shaping abiotic and biotic stress tolerance, plant architecture, seed oil content, and fiber development. We also suggest the application of novel technologies and strategies to facilitate genome-based crop breeding. Explosive growth in the amount of novel genomic data, identified genes, gene modules, and pathways is now enabling researchers to utilize multidisciplinary genomics-enabled breeding strategies to cultivate “super cotton”, synergistically improving multiple traits. These strategies must rise to meet urgent demands for a sustainable cotton industry.
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RNA-directed DNA methylation has an important developmental function in Arabidopsis that is masked by the chromatin remodeler PICKLE
Rong Yang, Li He, Huan Huang, Jian-Kang Zhu, Rosa Lozano-Duran and Heng Zhang
J Integr Plant Biol 2020, 62 (11): 1647-1652.  
doi: 10.1111/jipb.12979
Abstract (Browse 442)  |   Save
In Arabidopsis, RNA‐directed DNA methylation (RdDM) is required for the maintenance of CHH methylation, and for de novo methylation in all (CG, CHG, and CHH) contexts, but no obvious effect of RdDM deficiency on plant development has been found to date. We show that the combination of mutations in the chromatin remodeler PKL and RdDM components results in developmental alterations, which appear in a SUPPRESSOR OF DRM1 DRM2 CMT3 (SDC)‐dependent manner.
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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 438)  |   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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Cited: Web of Science(60)
  
MYB106 is a negative regulator and a substrate for CRL3BPM E3 ligase in regulating flowering time in Arabidopsis thaliana
Liu Hong, Fangfang Niu, Youshun Lin, Shuang Wang, Liyuan Chen and Liwen Jiang
J Integr Plant Biol 2021, 63 (6): 1104-1119.  
DOI: 10.1111/jipb.13071
Abstract (Browse 434)  |   Save
Flowering time is crucial for successful reproduction in plants, the onset and progression of which are strictly controlled. However, flowering time is a complex and environmentally responsive history trait and the underlying mechanisms still need to be fully characterized. Post-translational regulation of the activities of transcription factors (TFs) is a dynamic and essential mechanism for plant growth and development. CRL3BPM E3 ligase is a CULLIN3-based E3 ligase involved in orchestrating protein stability via the ubiquitin proteasome pathway. Our study shows that the mutation of MYB106 induced early flowering phenotype while over-expression of MYB106 delayed Arabidopsis flowering. Transcriptome analysis of myb106 mutants reveals 257 differentially expressed genes between wild type and myb106-1 mutants, including Flowering Locus T (FT) which is related to flowering time. Moreover, in vitro electrophoretic mobility shift assays (EMSA), in vivo chromatin immunoprecipitation quantitative polymerase chain reaction (ChIP-qPCR) assays and dual luciferase assays demonstrate that MYB106 directly binds to the promoter of FT to suppress its expression. Furthermore, we confirm that MYB106 interacts with BPM proteins which are further identified by CRL3BPM E3 ligases as the substrate. Taken together, we have identified MYB106 as a negative regulator in the control of flowering time and a new substrate for CRL3BPM E3 ligases in Arabidopsis.
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bZIP71 delays flowering by suppressing Ehd1 expression in rice
Xiufeng Li, Xiaojie Tian, Mingliang He, Xinxin Liu, Zhiyong Li, Jiaqi Tang, Enyang Mei, Min Xu, Yingxiang Liu, Zhenyu Wang, Qingjie Guan, Wei Meng, Jun Fang, Jian Zhang and Qingyun Bu
J Integr Plant Biol 2022, 64 (7): 1352-1363.  
DOI: 10.1111/jipb.13275
Abstract (Browse 432)  |   Save

Flowering time is a fundamental factor determining the global distribution and final yield of rice (Oryza sativa). Although diverse flowering time genes have been reported in this crop, the transcriptional regulation of its key flowering genes are poorly understood. Here, we report that a basic leucine zipper transcription factor, bZIP71, functions as a flowering repressor. The overexpression of bZIP71 delays flowering, while the bzip71 mutant flowers early in both long-day and short-day conditions. A genetic analysis showed that the regulation of flowering by bZIP71 might be independent of Heading date 2 (Hd2), Hd4, and Hd5. Importantly, bZIP71 directly associates with the Early heading date 1 (Ehd1) promoter and represses its transcription, and genetically the function of bZIP71 is impaired in the ehd1 mutant. Moreover, bZIP71 interacts with major components of polycomb repressive complex 2 (PRC2), SET domain group protein 711 (SDG711), and Fertilization independent endosperm 2 (FIE2), through which bZIP71 regulates the H3K27me3 level of Ehd1. Taken together, we present a transcriptional regulatory mechanism in which bZIP71 enhances the H3K27me3 level of Ehd1 and transcriptionally represses its expression, which not only offers a novel insight into a flowering pathway, but also provides a valuable putative target for the genetic engineering and breeding of elite rice cultivars.

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A rare Waxy allele coordinately improves rice eating and cooking quality and grain transparency
Changquan Zhang, Yong Yang, Shengjie Chen, Xueju Liu, Jihui Zhu, Lihui Zhou, Yan Lu, Qianfeng Li, Xiaolei Fan, Shuzhu Tang, Minghong Gu and Qiaoquan Liu
J Integr Plant Biol 2021, 63 (5): 889-901.  
doi: 10.1111/jipb.13010
Abstract (Browse 426)  |   Save
In rice (Oryza sativa), amylose content (AC) is the major factor that determines eating and cooking quality (ECQ). The diversity in AC is largely attributed to natural allelic variation at the Waxy (Wx) locus. Here we identified a rare Wx allele, Wxmw, which combines a favorable AC, improved ECQ and grain transparency. Based on a phylogenetic analysis of Wx genomic sequences from 370 rice accessions, we speculated that Wxmw may have derived from recombination between two important natural Wx alleles, Wxin and Wxb. We validated the effects of Wxmw on rice grain quality using both transgenic lines and near‐isogenic lines (NILs). When introgressed into the japonica Nipponbare (NIP) background, Wxmw resulted in a moderate AC that was intermediate between that of NILs carrying the Wxb allele and NILs with the Wxmp allele. Notably, mature grains of NILs fixed for Wxmw had an improved transparent endosperm relative to soft rice. Further, we introduced Wxmw into a high‐yielding japonica cultivar via molecular marker‐assisted selection: the introgressed lines exhibited clear improvements in ECQ and endosperm transparency. Our results suggest that Wxmw is a promising allele to improve grain quality, especially ECQ and grain transparency of high‐yielding japonica cultivars, in rice breeding programs.
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Cited: Web of Science(30)
  
Variations in OsSPL10 confer drought tolerance by directly regulating OsNAC2 expression and ROS production in rice
Yingxiu Li, Shichen Han, Xingming Sun, Najeeb Ullah Khan, Qun Zhong, Zhanying Zhang, Hongliang Zhang, Feng Ming, Zichao Li and Jinjie Li
J Integr Plant Biol 2023, 65 (4): 918-933.  
DOI: 10.1111/jipb.13414
Abstract (Browse 423)  |   Save
Drought is a major factor restricting the production of rice (Oryza sativa L.). The identification of natural variants for drought stress‐ related genes is an important step toward developing genetically improved rice varieties. Here, we characterized a member of the SQUAMOSA PROMOTER BINDING PROTEIN‐LIKE (SPL) family, OsSPL10, as a transcription factor involved in the regulation of drought tolerance in rice. OsSPL10 appears to play a vital role in drought tolerance by controlling reactive oxygen species (ROS) production and stomatal movements. Haplotype and allele frequency analyses of OsSPL10 indicated that most upland rice and improved lowland rice varieties harbor the OsSPL10Hap1 allele, whereas the OsSPL10Hap2 allele was mainly present in lowland and landrace rice varieties. Importantly, we demonstrated that the varieties with the OsSPL10Hap1 allele showed low expression levels of OsSPL10 and its downstream gene, OsNAC2, which decreases the expression of OsAP37 and increases the expression of OsCOX11, thus preventing ROS accumulation and programmed cell death (PCD). Furthermore, the knockdown or knockout of OsSPL10 induced fast stomatal closure and prevented water loss, thereby improving drought tolerance in rice. Based on these observations, we propose that OsSPL10 confers drought tolerance by regulating OsNAC2 expression and that OsSPL10Hap1 could be a valuable haplotype for the genetic improvement of drought tolerance in rice.
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Molecular mechanisms for the photoperiodic regulation of flowering in soybean
Xiaoya Lin, Baohui Liu, James L. Weller, Jun Abe and Fanjiang Kong
J Integr Plant Biol 2021, 63 (6): 981-994.  
doi: 10.1111/jipb.13021
Abstract (Browse 423)  |   Save
Photoperiodic flowering is one of the most important factors affecting regional adaptation and yield in soybean (Glycine max). Plant adaptation to long-day conditions at higher latitudes requires early flowering and a reduction or loss of photoperiod sensitivity; adaptation to short-day conditions at lower latitudes involves delayed flowering, which prolongs vegetative growth for maximum yield potential. Due to the influence of numerous major loci and quantitative trait loci (QTLs), soybean has broad adaptability across latitudes. Forward genetic approaches have uncovered the molecular basis for several of these major maturity genes and QTLs. Moreover, the molecular characterization of orthologs of Arabidopsis thaliana flowering genes has enriched our understanding of the photoperiodic flowering pathway in soybean. Building on early insights into the importance of the photoreceptor phytochrome A, several circadian clock components have been integrated into the genetic network controlling flowering in soybean: E1, a repressor of FLOWERING LOCUS T orthologs, plays a central role in this network. Here, we provide an overview of recent progress in elucidating photoperiodic flowering in soybean, how it contributes to our fundamental understanding of flowering time control, and how this information could be used for molecular design and breeding of high-yielding soybean cultivars.
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Cited: Web of Science(53)
  
The plant cell wall: Biosynthesis, construction, and functions
Baocai Zhang, Yihong Gao, Lanjun Zhang and Yihua Zhou
J Integr Plant Biol 2021, 63 (1): 251-272.  
doi: 10.1111/jipb.13055
Abstract (Browse 422)  |   Save
The plant cell wall is composed of multiple biopolymers, representing one of the most complex structural networks in nature. Hundreds of genes are involved in building such a natural masterpiece. However, the plant cell wall is the least understood cellular structure in plants. Due to great progress in plant functional genomics, many achievements have been made in uncovering cell wall biosynthesis, assembly, and architecture, as well as cell wall regulation and signaling. Such information has significantly advanced our understanding of the roles of the cell wall in many biological and physiological processes and has enhanced our utilization of cell wall materials. The use of cutting‐edge technologies such as single‐molecule imaging, nuclear magnetic resonance spectroscopy, and atomic force microscopy has provided much insight into the plant cell wall as an intricate nanoscale network, opening up unprecedented possibilities for cell wall research. In this review, we summarize the major advances made in understanding the cell wall in this era of functional genomics, including the latest findings on the biosynthesis, construction, and functions of the cell wall.
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Cited: Web of Science(63)
  
The nodulation and nyctinastic leaf movement is orchestrated by clock gene LHY in Medicago truncatula
Yiming Kong, Lu Han, Xiu Liu, Hongfeng Wang, Lizhu Wen, Xiaolin Yu, Xiaodong Xu, Fanjiang Kong, Chunxiang Fu, Kirankumar S. Mysore, Jiangqi Wen and Chuanen Zhou
J Integr Plant Biol 2020, 62 (12): 1880-1895.  
DOI: 10.1111/jipb.12999
Abstract (Browse 419)  |   Save

As sessile organisms, plants perceive, respond, and adapt to the environmental changes for optimal growth and survival. The plant growth and fitness are enhanced by circadian clocks through coordination of numerous biological events. In legume species, nitrogen‐fixing root nodules were developed as the plant organs specialized for symbiotic transfer of nitrogen between microsymbiont and host. Here, we report that the endogenous circadian rhythm in nodules is regulated by MtLHY in legume species Medicago truncatula. Loss of function of MtLHY leads to a reduction in the number of nodules formed, resulting in a diminished ability to assimilate nitrogen. The operation of the 24‐h rhythm in shoot is further influenced by the availability of nitrogen produced by the nodules, leading to the irregulated nyctinastic leaf movement and reduced biomass in mtlhy mutants. These data shed new light on the roles of MtLHY in the orchestration of circadian oscillator in nodules and shoots, which provides a mechanistic link between nodulation, nitrogen assimilation, and clock function.

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Comprehensive identification of lysine 2‐hydroxyisobutyrylated proteins in Ustilaginoidea virens reveals the involvement of lysine 2‐hydroxyisobutyrylation in fungal virulence
Xiaoyang Chen, Xiabing Li, Pingping Li, Xiaolin Chen, Hao Liu, Junbin Huang, Chaoxi Luo, Tom Hsiang and Lu Zheng
J Integr Plant Biol 2021, 63 (2): 409-425.  
doi: 10.1111/jipb.13066
Abstract (Browse 418)  |   Save
Lysine 2‐hydroxyisobutyrylation (Khib) is a newly identified post‐translational modification (PTM) that plays important roles in transcription and cell proliferation in eukaryotes. However, its function remains unknown in phytopathogenic fungi. Here, we performed a comprehensive assessment of Khib in the rice false smut fungus Ustilaginoidea virens, using Tandem Mass Tag (TMT)‐based quantitative proteomics approach. A total of 3 426 Khib sites were identified in 977 proteins, suggesting that Khib is a common and complex PTM in U. virens. Our data demonstrated that the 2‐hydroxyisobutyrylated proteins are involved in diverse biological processes. Network analysis of the modified proteins revealed a highly interconnected protein network that included many well‐studied virulence factors. We confirmed that the Zn‐binding reduced potassium dependency3‐type histone deacetylase (UvRpd3) is a major enzyme that removes 2‐hydroxyisobutyrylation and acetylation in U. virens. Notably, mutations of Khib sites in the mitogen‐activated protein kinase (MAPK) UvSlt2 significantly reduced fungal virulence and decreased the enzymatic activity of UvSlt2. Molecular dynamics simulations demonstrated that 2‐hydroxyisobutyrylation in UvSlt2 increased the hydrophobic solvent‐accessible surface area and thereby affected binding between the UvSlt2 enzyme and its substrates. Our findings thus establish Khib as a major post‐translational modification in U. virens and point to an important role for Khib in the virulence of this phytopathogenic fungus.
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Ethylene and salicylic acid synergistically accelerate leaf senescence in Arabidopsis
Chaoqi Wang, Shouyi Dai, Zhong‐Lin Zhang, Wenqing Lao, Ruiying Wang, Xianqing Meng and Xin Zhou
J Integr Plant Biol 2021, 63 (5): 828-833.  
doi: 10.1111/jipb.13075
Abstract (Browse 414)  |   Save
The phytohormones ethylene and salicylic acid (SA) have long been known to promote senescence, but their interplay during this process remains elusive. Here we report the synergistic effects of ethylene and SA on promoting leaf senescence in Arabidopsis. EIN3, a key transcription factor of ethylene signaling, physically interacted with the core SA signaling regulator NPR1 in senescing leaves. EIN3 and NPR1 synergistically promoted the expression of the senescence‐associated genes ORE1 and SAG29. The senescence phenotype was more delayed for the ein3eil1npr1 triple mutant than ein3eil1 or npr1 with ethylene or/and SA treatment. NPR1‐promoted leaf senescence may depend on functional EIN3/EIL1.
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