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Authors: Yongguo Xue, Yuntong Zhang, Jinming Shan, Yujia Ji, Xiaoming Zhang, Wenbin Li, Dongmei Li and Lin Zhao.
International Journal of Molecular Sciences (2022)
Abstract: "Plant height is an important component of plant architecture, and significantly affects crop quality and yield. A soybean GmRAV (Related to ABI3/VP1) transcription factor containing both AP2 and B3 domains is a growth repressor. Three GmRAV-overexpressing (GmRAV-ox) transgenic lines displayed extremely shorter height and shortened internodes compared with control plants, whereas transgenic inhibition of GmRAV expression resulted in increased plant height. GmRAV-ox soybean plants showed a low active gibberellin level and the dwarf phenotype could be rescued by treatment with exogenous GA3 treatment. ChIP (Chromatin immunoprecipitation)-qPCR assay showed that GmRAV could directly regulate the expression of the GA4 biosynthetic genes GA3-oxidase (GmGA3ox) by binding two CAACA motifs in the GmGA3ox promoter. The GmGA3ox promoter was bound by GmRAV, whose expression levels in leaves were both elevated in GmRAV-i-3 and decreased in GmRAV-ox-7 soybean plants. Transient expression assay in N. benthamiana also showed that the proGmRAV:GmRAV-3F6H effector strongly repressed the expression of LUC reporter gene driven by GmGA3ox promoter containing two CAACA motifs. Together, our results suggest that GmRAV protein repressed the expression of GmGA3ox by directly binding to the two CAACA motifs in the promoter to limit soybean plant height."
Authors: Hongchun Xiong, Chunyun Zhou, Meiyu Fu, Huijun Guo, Yongdun Xie, Linshu Zhao, Jiayu Gu, Shirong Zhao, Yuping Ding, Yuting Li, Jiazi Zhang, Ke Wang, Xuejun Li and Luxiang Liu.
Molecular Plant (2022)
Excerpts: "..continuous efforts have been made to search for “Green Revolution” replacement genes in wheat. Rht8 is a semi-dwarfing gene that complements well the shortcomings of Rht-B1b and Rht-D1b, thus has been widely introgressed into wheat varieties across the globe...."
"Rht8 plants are gibberellic acid (GA) sensitive (Ellis et al., 2005). Consistently, we showed that treatment with exogenous GA3 could effectively restore the semi-dwarf phenotype of Rht8-2 and Rht8-3 (Figure 1F, Supplemental Figure 8), hinting a role of Rht8 in regulating GA biosynthesis. We further demonstrated that GA biosynthetic genes, such as GA20ox-2 and GA13ox, were significantly down-regulated in both 66 mutants (Figure 1G, Supplemental Figure 9)."
"To sum up, the present work identifies two new semi-dwarf wheat mutants that are allelic to Rht8, reveals the complexity and evolutionarily history of Rht8 in common wheat. Rht8 encodes a protein containing a Zinc finger BED-type motif and an RNase H-like domain that regulates plant height via influencing bioactive GA biosynthesis."
Authors: Jie Liu, Yingyin Yao, Mingming Xin, Huiru Peng, Zhongfu Ni and Qixin Sun.
Journal of Integrative Plant Biology (2022)
Abstract: "Bread wheat (Triticum aestivum L., AABBDD, 2n = 6x = 42), which accounts for most of the cultivated wheat crop worldwide, is a typical allohexaploid with a genome derived from three diploid wild ancestors. Bread wheat arose and evolved via two sequential allopolyploidization events and was further polished through multiple steps of domestication. Today cultivated allohexaploid bread wheat has numerous advantageous traits, including adaptive plasticity, favorable yield traits, and extended end-use quality, that have enabled its cultivation well beyond the ranges of its tetraploid and diploid progenitors to become a global staple food crop. In the past decade, rapid advances in wheat genomic research have considerably accelerated our understanding of the bases for the shaping of complex agronomic traits in this polyploid crop. Here, we summarize recent advances in characterizing major genetic factors underlying the origin, evolution, and improvement of polyploid wheats. We end with a brief discussion of the future prospects for the design of gene cloning strategies and modern wheat breeding."
Authors: Jiaqi Zhang, Xiuying Gao, Guang Cai, Yuji Wang, Jianbo Li, Huaying Du, Ruqin Wang, Hongsheng Zhang and Ji Huang.
Rice (2021)
Abstract: "Background - Grain size is one of the major determinants of cereal crop yield. As a class of plant polyhydroxysteroids, brassinosteroids (BRs) play essential roles in the regulation of grain size and plant architecture in rice. In a previous research, we cloned qGL3/OsPPKL1 encoding a protein phosphatase with Kelch-like repeat domains, which negatively regulates BR signaling and grain length in rice. Results - Here, we screened qGL3-interacting proteins (GIPs) via yeast two-hybrid assay and analyzed the phenotypes of the T-DNA insertion mutants of GIPs. Among these mutants, mutant osak3 presents shorter grain length and dwarfing phenotype. OsAK3 encodes an adenylate kinase, which regulates grain size by controlling cell expansion of rice spikelet glume. Overexpression of OsAK3 resulted in longer grain length. OsAK3 interacts with qGL3 in vivo and in vitro. Lamina inclination, coleoptile elongation and root inhibition experiments showed that the osak3 mutant was less sensitive to exogenous brassinolide (BL) treatment. The transcriptional level of OsAK3 was up-regulated under BL induction. In addition, RNA-Seq data indicate that OsAK3 is involved in a variety of biological processes that regulate BR signaling and grain development in rice. Conclusions - Our study reveals a novel BR signaling component OsAK3 in the regulation of grain length, and provides novel clues for uncovering the potential functions of OsAK3 in rice growth and development."
Authors: Jiajun Liu, Xi’an Shi, Zhongyuan Chang, Yanfeng Ding and Chengqiang Ding.
Plant and Cell Physiology (2022)
Abstract: "The essential role of auxin in plant growth and development is well known. Pathways related to auxin synthesis, transport, and signaling have been extensively studied in recent years, and the PIN-FORMED (PIN) protein family has been identified to be pivotal for polar auxin transport (PAT) and distribution. However, research focused on the functional characterization of PIN proteins in rice is still lacking. In this study, we investigated the expression and function of OsPIN1c and OsPIN1d in the japonica rice variety (Nipponbare) using gene knockout and high-throughput RNA sequencing (RNA-Seq) analysis. Results showed that OsPIN1c and OsPIN1d were mainly expressed in young panicles and exhibited a redundant function. Furthermore, OsPIN1c or OsPIN1d loss-of-function mutants presented a mild phenotype compared with the wild type. In addition to significantly decreased plant height and tiller number, panicle development was severely disrupted in double mutant lines of OsPIN1c and OsPIN1d. Severe defects included smaller inflorescence meristem (IM) and panicle sizes, fewer primary branches, elongated bract leaves, non-degraded hair, and no spikelet growth. Interestingly, ospin1cd-3, a double mutant line with functional retention of OsPIN1d, showed milder defects than those observed in other mutants. Additionally, several critical regulators of reproductive development, such as OsPID, LAX1, OsMADS1, and OsSPL14/IPA1, were differentially expressed in ospin1c-1 ospin1d-1, supporting the hypothesis that OsPIN1c and OsPIN1d are involved in regulating panicle development. Therefore, this study provides novel insights into the auxin pathways that regulate plant reproductive development in monocots."
Authors: Ruijuan Yang, Zhenying Wu, Chen Bai, Zhichao Sun, Mengqi Wang, Yuzhu Huo, Hailing Zhang, Yamei Wang, Huapeng Zhou, Shaojun Dai, Wenwen Liu and Chunxiang Fu.
Horticulture Research (2021)
Abstract: "Switchgrass (Panacum virgatum L.) is an important perennial, noninvasive, tall ornamental grass that adds color and texture to gardens and landscapes. Moreover, switchgrass has been considered a forage and bioenergy crop because of its vigorous growth, low-input requirements, and broad geography. Here, we identified PvWOX3a from switchgrass, which encodes a WUSCHEL-related homeobox transcription factor. Transgenic overexpression of PvWOX3a in switchgrass increased stem length, internode diameter, and leaf blade length and width, all of which contributed to a 95% average increase in dry weight biomass compared with control plants. Yeast one-hybrid and transient dual-luciferase assays showed that PvWOX3a can repress the expression of gibberellin 2-oxidase and cytokinin oxidase/dehydrogenase through apparently direct interaction with their promoter sequences. These results suggested that overexpression of PvWOX3a could increase gibberellin and cytokinin levels in transgenic switchgrass plants, which promotes cell division, elongation, and vascular bundle development. We also overexpressed PvWOX3a in a transgenic miR156-overexpressing switchgrass line that characteristically exhibited more tillers, thinner internodes, and narrower leaf blades. Double transgenic switchgrass plants displayed significant increases in internode length and diameter, leaf blade width, and plant height but retained a tiller number comparable to that of plants expressing miR156 alone. Ultimately, the double transgenic switchgrass plants produced 174% more dry-weight biomass and 162% more solubilized sugars on average than control plants. These findings indicated that PvWOX3a is a viable potential genetic target for engineering improved shoot architecture and biomass yield of horticulture, fodder, and biofuel crops."
Authors: Shuchao Dong, Danuse Tarkowska, Mastoureh Sedaghatmehr, Maryna Welsch, Saurabh Gupta, Bernd Mueller-Roeber and Salma Balazadeh.
Molecular Plant (2022)
Abstract: "The phytohormones gibberellins (GAs) play fundamental roles in almost every aspect of plant growth and development. Although there is good knowledge about GA biosynthetic and signaling pathways, factors contributing to the mechanisms homeostatically controlling GA levels remain largely unclear. Here, we demonstrate that homeobox transcription factor HB40 of the HD-Zip family in Arabidopsis thaliana regulates GA content at two additive control levels. We show that HB40 expression is induced by GA and in turn reduces the levels of endogenous bioactive GAs by a simultaneous reduction of GA biosynthesis and increased GA deactivation. Hence, HB40 overexpression leads to typical GA-deficiency traits, such as small rosettes, reduced plant height, delayed flowering, and male sterility. In contrast, a loss-of-function hb40 mutation enhances GA-controlled growth. Genome-wide RNA-sequencing combined with molecular-genetic analyses revealed that HB40 directly activates transcription of JUNGBRUNNEN1 (JUB1), a key TF repressing growth by suppressing GA biosynthesis and signaling. HB40 also activates genes encoding GA 2-oxidases (GA2oxs) which are major GA catabolic enzymes. The effect of HB40 is ultimately mediated through induction of nuclear growth-repressing DELLA proteins. Our results thus uncover an important role of the HB40/JUB1/GA2ox/DELLA regulatory network in controlling GA homeostasis during plant growth."
Authors: Fengxia Wang, Zipeng Yu, Maolin Zhang, Mengli Wang, Xiaoduo Lu, Xia Liu, Yubin Li ,Xiansheng Zhang, Bao-cai Tan, Cuiling Li and Zhaojun Ding.
Plant Biotechnology Journal (2021)
Abstract: "Maize height is determined by the number of nodes and the length of internodes. Node number is driven by intercalary meristem formation and internode length by intercalary cell elongation respectively. However, mechanisms regulating establishment of nodes and internode growth is unclear. We screened EMS-induced maize mutants and identified a dwarf mutant zm66, linked to a single base change in TERMINAL EAR 1 (ZmTE1). Detailed phenotypic analysis revealed that zm66 (zmte1-2) has shorter internodes and increased node numbers, caused by decreased cell elongation and disordered intercalary meristem formation, respectively. Transcriptome analysis showed that auxin signaling genes are also dysregulated in zmte1-2, as are cell elongation and cell cycle-related genes. This argues that ZmTE1 regulates auxin signaling, cell division and cell elongation. We found that the ZmWEE1 kinase phosphorylates ZmTE1, thus confining it to the nucleus and probably reducing cell division. In contrast, the ZmPP2Ac-2 phosphatase promotes dephosphorylation and cytoplasmic localization of ZmTE1, as well as cell division. Taken together, ZmTE1, a key regulator of plant height, is responsible for maintaining organized formation of internode meristems and rapid cell elongation. ZmWEE1 and ZmPP2Ac-2 might balance ZmTE1 activity, controlling cell division and elongation to maintain normal maize growth."
Authors: Wei Dong, Dewei Wu, Caihui Wang, Ying Liu and Defeng Wu.
Plant Science (2021)
Highlights • The dsh is a GA-deficient mutant and useful germplasm resource. • Mutations in the promoter of ClaGA20ox2 mediate the dwarfism.
Abstract: "Developing dwarf watermelon is a major objective among breeders. The dsh dwarf watermelon germplasm developed in our laboratory is genetically stable. We previously produced preliminary evidence that Cla010726, which encodes a gibberellin 20-oxidase-like protein, is the primary gene controlling dwarfism in watermelon. However, the underlying genetic mechanism was unknown. In this study, we characterized the spontaneous recessive mutant dsh, which is a gibberellin (GA)-deficient mutant. Many of the phenotypic traits of dsh plants are similar to those of known GA-deficient mutants. The dsh plants were sensitive to exogenous bioactive GAs, which increased seedling height. Moreover, a quantitative analysis of endogenous GA3 proved that the bioactive GA3 content was substantially lower than normal in dsh. Additionally, the T5 ClaGA20ox RNAi plants generally exhibited dwarfism, with short stems and internodes as well as small leaves and fruit. An examination of the transgenic plants carrying the ClaGA20ox1 promoter-GUS and mutant ClaGA20ox2 promoter-GUS constructs confirmed that two promoter sites are involved in the regulation of ClaGA20ox expression. Hence, mutations in the promoter of the GA20ox gene, which encodes a key enzyme involved in gibberellin biosynthesis, lead to the dwarfism of watermelon plants. The dsh mutant is a potentially useful germplasm resource for developing new watermelon varieties exhibiting dwarfism."
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Authors: Daniela Gómez-Soto, Isabel Allona and Mariano Perales.
Frontiers in Plant Science (2022)
Abstract: "The adaptation and survival of boreal and temperate perennials relies on the precise demarcation of the growing season. Seasonal growth and development are defined by day length and temperature signals. Under long-day conditions in spring, poplar FLOWERING LOCUS T2 (FT2) systemically induces shoot growth. In contrast, FT2 downregulation induced by autumnal short days triggers growth cessation and bud set. However, the molecular role of FT2 in local and long-range signaling is not entirely understood. In this study, the CRISPR/Cas9 editing tool was used to generate FT2 loss of function lines of hybrid poplar. Results indicate that FT2 is essential to promote shoot apex development and restrict internode elongation under conditions of long days. The application of bioactive gibberellins (GAs) to apical buds in FT2 loss of function lines was able to rescue bud set. Expression analysis of GA sensing and metabolic genes and hormone quantification revealed that FT2 boosts the 13-hydroxylation branch of the GA biosynthesis pathway in the shoot apex. Paclobutrazol treatment of WT leaves led to limited internode growth in the stem elongation zone. In mature leaves, FT2 was found to control the GA 13-hydroxylation pathway by increasing GA2ox1 and reducing GA3ox2 expression, causing reduced GA1 levels. We here show that in poplar, the FT2 signal promotes shoot apex development and restricts internode elongation through the GA 13-hydroxylation pathway."
Authors: Ming Li, Hui Zhang, Dongli He, Rebecca Njeri Damaris and Pingfang Yang.
Journal of Experimental Botany (2022)
Abstract: "Gibberellic acid (GA) is a vital phytohormone for plant growth and development. GA biosynthesis is a complex pathway regulated by various transcription factors. Here we report a stress-associated protein OsSAP8 negatively involved in GA biosynthesis. Overexpression of OsSAP8 in rice resulted in a semi-dwarfism phenotype and reduced endogenous GA3 content. By contrast, higher endogenous GA3 content and slightly higher plant height were observed in OsSAP8 knockout mutant. Subcellular localization analysis of OsSAP8 showed it could enter into nucleus. Based on electrophoretic mobility shift assay and yeast one hybrid experiment, OsSAP8 could bind to cis-acting regulatory element GADOWNAT of ent-kaurene oxidases (KO2, KO3, KO5). The results from dual-luciferase reporter assays showed OsSAP8 doesn’t activate LUC reporter gene expression. However, it could interact with OsbZIP58, which has strong transcriptional activation potential on OsKO2. Moreover, the interaction between OsSAP8, OsLOL1, and OsbZIP58 could reduce the promotive effect of transcription factor OsbZIP58 on OsKO2. These results provide some new clues on the regulation of GA biosynthesis in rice."
Authors: Lihua Zheng, Jiangqi Wen, Jinling Liu, Xiangzhao Meng, Peng Liu, Na Cao, Jiangli Dong and Tao Wan.
The Crop Journal (2022)
Abstract: "Alfalfa (Medicago sativa L.) is a nutritious forage crop with wide ecological adaptability. The molecular breeding of alfalfa is restricted by its heterozygous tetraploid genome and the difficult genetic manipulation process. Under time and resource constraints, we applied a more convenient approach. We investigated two MtGA3ox genes, MtGA3ox1 and MtGA3ox2, of Medicago truncatula, a diploid legume model species, finding that MtGA3ox1 plays a major role in GA-regulated plant architecture. Mutation of neither gene affected nitrogenase activity. These results suggest that MtGA3ox1 can be used in semidwarf and prostrate alfalfa breeding. Based on the M. truncatula MtGA3ox1 sequence, MsGA3ox1 was cloned from alfalfa, and two knockout targets were designed. An efficient CRISPR/Cas9-based genome editing protocol was used to generate msga3ox1 mutants in alfalfa. We obtained three lines that carried mutations in all four alleles in the T0 generation. Fifteen clonal plants were vegetatively propagated from each transgenic line using shoot cuttings. The plant height and internode length of msga3ox1 null mutants were significantly decreased. The number of total lateral branches, leaf/stem ratio and crude protein content of aerial plant parts of msga3ox1 mutants were significantly increased. Thus, we obtained semi-dwarf and prostrate alfalfa by gene editing."
Authors: Yuya Kumagai, Yuelin Liu, Haruyasu Hamada, Weifeng Luo, Jianghui Zhu, Misa Kuroki, Yozo Nagira, Naoaki Taoka, Etsuko Katoh and Ryozo Imai.
Plant Physiology (2022)
One-sentence summary: "Direct delivery of CRISPR/Cas9 ribonucleoproteins into the shoot apical meristem via particle bombardment enabled introduction of a semidwarf1-orthologous mutation into an elite wheat variety.
Excerpts: "These results suggest that GA20 oxidases within a clade have an evolutionarily conserved function. Thus, we concluded that TaSD-A1, TaSD-B1 and TaSD-D1 were the three wheat orthologs that are homologous to rice sd1. To create a tasd1 triple knockout mutant using CRISPR/Cas9 RNP, three single-guide RNA (sgRNA) target sequences (target_1, target_2, and target_3) were designed that commonly appear within the TaSD-A1, TaSD-B1, and TaSD-D1 genes (Fig. 2a)."
"The phenotype of the tasd1 mutant was analyzed in the E3 generation of the H7-1 line. Both wild-type (WT) and H7-1 mutant plants were grown under long day conditions in an environmentally-controlled growth room. The mutant plants exhibited greener leaf color and shorter plant height. The average final height of the plants was approximately 10% lower in the tasd1 mutant (Fig 2f), relative to the WT. The average total number of grains and grain weight were nearly equivalent in WT and tasd1 plants (Fig. 2g)."
Authors: Yanan Niu, Tianxiao Chen, Chenchen Zhao and Meixue Zhou.
Agronomy (2021)
Abstract: "Crop height not only determines plant resistance to lodging and crowding, but also affects crop architecture, apical dominance, biomass, and mechanical harvesting. Plant height is determined by the internode elongation, regulated by genes involved in gibberellin (GA) and brassinosteroid (BR) biosynthesis or related signaling networks. Plants’ genetic inability to synthesize or respond to GAs and BRs induce dwarfness. However, the signaling mechanisms of GAs and BRs for controlling plant height individually or collectively are still unclear. Since stem mechanically supports plant during the whole life span, components that affect stem physical strength are also important to crop lodging resistance. One of the major components is lignin, which forms stem structure, thus contributing to crop lodging resistance. In this review, we looked into the reported genes involved in lignin, GAs, and BRs biosynthesis and summarized the signaling networks centered by these genes. Then, we filled the knowledge gap by modifying plant height through interrupting normal GA and BR metabolism utilizing core gene inhibitors. Therefore, we highly endorsed the current approaches of using plant growth regulators (PRGs) to maintain an ideal plant height under lodging stress, and proposed possibilities of modifying crop culm strength against lodging as well."
Authors: Yaping Zhang, Ayla Norris, Michael S. Reid and Cai-Zhong Jiang.
Ornamental Plant Research (2021)
Abstract: "Improving plants' ability to survive under drought is of great importance to the horticultural industry. The plant hormone gibberellic acid (GA) mediates diverse aspects of plant growth and development. The Arabidopsis gibberellin acid insensitive mutant gai-1 displays reduced plant height, altered GA response, and enhanced drought resistance. However, overexpression of gai-1 using the constitutive 35S promoter would result in dwarf plants with drought resistance. Here, we tested the hypothesis that the temporary inhibition of cell growth caused by inducible expression of the gai-1 gene would lead to better drought resistance and improve crop productivity without an undesirable dwarf phenotype. We generated transgenic plants in which the gai-1 gene was over-expressed in petunia, under a stress-inducible RD29A promoter from Arabidopsis. When these plants were subjected to limited irrigation and drought treatments, transgenic plants showed phenotypes of darker green leaves and compact flowers compared to the wild type plants. Importantly, these transgenic plants recovered sooner than wild type and the empty vector-transformed control plants. This study provides evidence that temporary inhibition of cell growth caused by over-expression of the gai-1 mutant gene with a drought stress-inducible promoter leads to better drought resistance when the plants experience drought conditions".
Authors: Hongsheng Gao, Huiqing Huang, Kaifeng Lu, Cuiting Wang, Xiaohua Liu, Zhizhong Song, Houjun Zhou, Lei Yang, Bei Li, Chunyan Yu and Hongxia Zhang.
Plant Physiology and Biochemistry (2021)
Highlights: • Constitutive expression of OsCYP714D1 promoted growth and salt tolerance in transgenic poplar. • Expression of gibberellin synthesis and metabolism genes was affected in OsCYP714D1 transgenic poplar. • OsCYP714D1 transgenic poplar showed altered ion homeostasis and related gene expression.
Abstract: "Cytochrome P450 monooxygenases (CYP450s) play crucial roles in the regulation of plant growth and response to abiotic stress. However, their functions in woody trees are still largely unknown. Previously, we reported that expression of the rice cytochrome P450 monooxygenase gene OsCYP714D1 increased gibberellic acid (GA) accumulation and shoot growth in transgenic poplar. In this work, we demonstrate that expression of OsCYP714D1 improved the salt tolerance of transgenic poplar plants. Compared to wild type, plant height and K+ content were significantly higher, whereas plant growth inhibition and Na+ content were significantly lower, in transgenic plants grown under high salt stress condition. Transcriptomic analyses revealed that OsCYP714D1 expression up-regulated the expressions of GA biosynthesis, signaling and stress responsive genes in transgenic plants under both normal and high salt stress conditions. Further gene ontology (GO) analyses indicated that genes involved in plant hormone and ion metabolic activities were significantly enriched in transgenic plants. Our findings imply that OsCYP714D1 participated in the regulation of both shoot growth and salt resistance through regulating gibberellin and ion homeostasis in transgenic poplar, and it can be used as a candidate gene for the engineering of new tree varieties with improved biomass production and salt stress resistance."
Authors: Xuemei Si, Wanxin Wang, Ke Wang, Yunchuan Liu, Jiangping Bai, Yaxiong Meng, Xueyong Zhang and Hongxia Liu.
International Journal of Molecular Sciences (2021)
Abstract: "The elongation and development of wheat (Triticum aestivum L.) stem play an important role in plant architecture. The shortened stem would result in a sheathed spike and a low yield in crops. Unraveling the molecular mechanisms underlying a sheathed spike would be beneficial for plant architecture and yield improvement. We identified a novel gene, TaWUS-like (WUSCHEL-related homeobox-like), which regulated sheathed spike and plant architecture in wheat. The plant height of overexpression transgenic lines was significantly decreased and the spike was not completely elongated and enclosed in flag leaf sheaths. Moreover, the increase in tiller angle resulted in loose plant architecture and lower yield. The statistical and cytological analysis demonstrated that the length of the uppermost and secondary internode was significantly shortened, especially the uppermost internode which was only half the length of the wild-type. The size of parenchyma cells was obviously reduced and cell length on the longitudinal section was elongated insufficiently compared with wild-type. The analysis of hormone content showed that there was a lack of gibberellin A 3 (GA3) in internodes but a higher brassinosteroid (BR) content. TaWUS-like may inhibit the synthesis of GA3 and/or BR, thus affecting the function of signal transduction of these hormones, which further caused stem shortening and plant dwarfing in wheat."
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