Seed Biology

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    UBA2A regulates seed dormancy and the stability of chromatin-retained DOG1 messenger RNA
    Ce Wang, Lien Brzezniak, Sebastian Sacharowski, Michal Krzyszton, Veena Halale Manjunath, Mateusz Jan Olechowski, Anna Kulik, Szymon Swiezewski
    J Integr Plant Biol 2025, 67 (12): 3109-3122.  
    doi: 10.1111/jipb.70056
    Abstract (Browse 261)  |   Save
    Multiple factors control primary seed dormancy established during seed maturation and secondary seed dormancy initiated when a non-dormant imbibed seed is exposed to adverse conditions. A key player in the control of primary and secondary dormancy in Arabidopsis thaliana is the Delay of Germination 1 (DOG1) gene, the expression of which is extensively regulated at the transcriptional and co-transcriptional levels. Despite its importance, the influence of post-transcriptional messenger RNA (mRNA) processing and mRNA storage of DOG1 on the determination of dormancy depth remains elusive. Here, we show that the UBA2A protein, a member of the heterogeneous nuclear ribonucleoprotein (hnRNP) family, negatively regulates primary and secondary seed dormancy through the regulation of the DOG1 gene expression at the post-transcriptional level. uba2a mutants show higher levels of the DOG1 mRNA. Surprisingly, DOG1 gene transcription is not affected, as demonstrated by single-molecule fluorescent in situ hybridization, chromatin-attached mRNA analysis and Pol II chromatin immunoprecipitation (ChIP). Instead, our results show that the UBA2A protein decreases the stability of both chromatin-bound and cytoplasmic DOG1 mRNA pools, and results in higher chromatin retention of DOG1 mRNA in the uba2a mutant. Our study highlights chromatin retention and mRNA stability as important features of DOG1 gene expression regulation with a profound impact on dormancy establishment and shows that UBA2A protein, like its human homolog hnRNPAB, is most likely implicated in mRNA transport in the cell.
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    In Arabidopsis, expression of DELAY OF GERMINATION 1 (DOG1), a key regulator of seed dormancy, is regulated not only at the level of transcription but also post-transcriptionally, at the level of mRNA stability.
      
    Genetic optimization of the source, sink and flow for increasing seed oil content in rapeseed
    Wenhao Shen, Liangqian Yu, Qian Qu, Xu Han, Wei Ma, Feng Zu, Liang Guo, Shan Tang
    J Integr Plant Biol 2025, 67 (11): 2799-2815.  
    DOI: 10.1111/jipb.70017
    Abstract (Browse 424)  |   Save
    Rapeseed (Brassica napus) is one of the most important oilseed crops worldwide, with its seed oil content (SOC) and quality directly determining its economic value. To resolve the challenges of growing demand for vegetable oil and advancements in rapeseed production, substantial progress has been achieved in the genetic improvement of SOC. This review systemizes genetic optimization strategies across three hierarchical processes: source expansion via enhanced photosynthesis, optimized carbon allocation, and metabolic redirection of photoassimilates; sink enhancement through targeted elevation of fatty acid (FA) synthesis, triacylglycerol (TAG) assembly, and seed coat development coupled with suppression of lipolytic pathways; flow optimization by modifying carbon partitioning, sucrose phloem loading and channeling to developing seeds. We synthesize reported genetic determinants of these processes and underscore their potential for enhancing SOC. Furthermore, we postulate that synergistic integration of source–flow–sink coordination with push–pull–package–protect frameworks could maximize oil accumulation, thereby establishing a multi-tiered roadmap for transcending SOC ceilings in rapeseed.
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    This review synthesizes genetic determinants for boosting seed oil content in rapeseed and systematizes these across three processes: source expansion via boosted photosynthesis and carbon allocation; sink enhancement via elevated fatty acid synthesis and triacylglycerol assembly with suppressed lipolysis; and flow optimization via modified carbon partitioning and sucrose transport.
      
    A major latex protein, GsMLP328, modulates seed traits in soybean
    Zhongyi Yang, Shaoqi Lu, Wei Li, Zhen Wang, Dezhou Hu, Xiaoyue Su, Fei Wu, Shupeng Dong, Xuan Cui, Yu Zhang, Yidan Zhang, Xiao Li, Qingxin Song, Deyue Yu, Guizhen Kan, Fang Huang
    J Integr Plant Biol 2025, 67 (11): 2790-2792.  
    doi: 10.1111/jipb.70018
    Abstract (Browse 310)  |   Save
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    The Major Latex Protein GsMLP328 modulates seed weight and protein content in soybean.
      
    Temperature-dependent embryoless seeds improve milled rice yield and storability
    Nai-Qian Dong, Hong-Xuan Lin
    J Integr Plant Biol 2025, 67 (11): 2784-2786.  
    doi: 10.1111/jipb.70028
    Abstract (Browse 217)  |   Save
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    During rice seed storage, lipid hydrolysis and oxidation in the embryo generate off-flavors. This Commentary examines a study by Wang et al., who demonstrated that manipulating OsBZR4 in various rice cultivars induces a high proportion of embryoless seeds by altering auxin levels and spatial distribution during early embryogenesis—a process intensified under elevated temperatures.
      
    FGM1/rPPR4-dependent female gamete maturation is essential for seed development initiation in Arabidopsis
    Feng Gong, Rongxin Lin, Zonglin Liu, Xiaorong Huang, Meng‐xiang Sun, Xiongbo Peng
    J Integr Plant Biol 2025, 67 (8): 2214-2228.  
    DOI: 10.1111/jipb.13922
    Abstract (Browse 335)  |   Save
    Gamete maturation is critical for fertility of both animals and plants; however, the molecular mechanisms underlying these processes remain poorly understood in plants. Here, we report the Female Gamete Maturation 1 (FGM1/rPPR4), a component of mitoribosome large subunit in Arabidopsis directly interacts with the mitochondrial protein GAMETE CELL DEFECTIVE 1 (GCD1) and plays an essential role in female gamete maturation and subsequent zygote–embryo transition and endosperm development. We reveal that FGM1/rPPR4, assisted by GCD1, is an essential factor for female gamete maturation. We also confirm that female gamete maturation is necessary for the capacity of post-fertilization zygote–embryo transition and endosperm development, but not for double fertilization, indicating that essential mechanisms are established during female gamete maturation to provide a molecular basis for seed development initiation and plant fertility.
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    FEMALE GAMETE MATURATION 1/PENTATRICOPEPTIDE REPEAT PROTEIN 4), a component of the mitochondrial ribosome large subunit, directly interacts with another mitochondrial protein GAMETE CELL DEFECTIVE 1 to play an essential role in maturation of egg cell and central cell. The gamete maturation is critical for post-fertilization embryo and endosperm development.
      
    PSC1, a basic/helix–loop–helix transcription factor controlling the purplish-red testa trait in peanut
    Kunkun Zhao, Jingjing Zhang, Yi Fan, Xufa Du, Shuliang Zhu, Zhongfeng Li, Ding Qiu, Zenghui Cao, Qian Ma, Yaoyao Li, Di Cao, Sasa Hu, Kai Zhao, Fangping Gong, Rui Ren, Xingli Ma, Xingguo Zhang, Dongmei Yin
    J Integr Plant Biol 2025, 67 (5): 1364-1378.  
    DOI: 10.1111/jipb.13847
    Abstract (Browse 404)  |   Save
    Seed color is a key agronomic trait in crops such as peanut, where it is a vital indicator of both nutritional and commercial value. In recent years, peanuts with darker seed coats have gained market attention due to their high anthocyanin content. Here, we used bulk segregant analysis to identify the gene associated with the purplish-red coat trait and identified a novel gene encoding a basic/helix–loop–helix transcription factor, PURPLE RED SEED COAT1 (PSC1), which regulates the accumulation of anthocyanins in the seed coat. Specifically, we found that a 35-bp insertion in the PSC1 promoter increased the abundance of PSC1 mRNA. Transcriptomic and metabolomic analyses indicated that the purplish-red color of the seed coat was the result of decreased expression of anthocyanidin reductase (ANR), leading to increased accumulation of delphinidin, cyanidin, and pelargonidin derivatives. Further analysis revealed that PSC1 interacts with AhMYB7 to form a complex that specifically binds to the ANR promoter to suppress its expression, resulting in increased anthocyanin accumulation. Moreover, overexpression of PSC1 increased anthocyanin content in Arabidopsis thaliana and peanut callus. Our study reveals a new gene that controls seed coat color by regulating anthocyanin metabolism and provides a valuable genetic resource for breeding peanuts with a purplish-red seed coat.
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    The basic/helix-loop-helix transcription factor PURPLE RED SEED COAT1 regulates the accumulation of anthocyanins in the seed coat in peanut and interacts with the MYB transcription factor AhMYB7 to form a complex binding to the promoter of the anthocyanidin reductase gene ANR to suppress its expression, resulting in increased anthocyanin accumulation.
      
    The comprehensive regulatory network in seed oil biosynthesis
    Wei Wei, Long-Fei Wang, Jian-Jun Tao, Wan-Ke Zhang, Shou-Yi Chen, Qingxin Song, Jin-Song Zhang
    J Integr Plant Biol 2025, 67 (3): 649-668.  
    doi: 10.1111/jipb.13834
    Abstract (Browse 373)  |   Save
    Plant oils play a crucial role in human nutrition, industrial applications and biofuel production. While the enzymes involved in fatty acid (FA) biosynthesis are well-studied, the regulatory networks governing these processes remain largely unexplored. This review explores the intricate regulatory networks modulating seed oil biosynthesis, focusing on key pathways and factors. Seed oil content is determined by the efficiency of de novo FA synthesis as well as influenced by sugar transport, lipid metabolism, FA synthesis inhibitors and fine-tuning mechanisms. At the center of this regulatory network is WRINKLED1 (WRI1), which plays a conserved role in promoting seed oil content across various plant species. WRI1 interacts with multiple proteins, and its expression level is regulated by upstream regulators, including members of the LAFL network. Beyond the LAFL network, we also discuss a potential nuclear factor-Y (NF-Y) regulatory network in soybean with an emphasis on NF-YA and NF-YB and their associated proteins. This NF-Y network represents a promising avenue for future efforts aimed at enhancing oil accumulation and improving stress tolerance in soybean. Additionally, the application of omics-based approaches is of great significance. Advances in omics technologies have greatly facilitated the identification of gene resources, opening new opportunities for genetic improvement. Importantly, several transcription factors involved in oil biosynthesis also participate in stress responses, highlighting a potential link between the two processes. This comprehensive review elucidates the complex mechanisms underlying the regulation of oil biosynthesis, offering insights into potential biotechnological strategies for improving oil production and stress tolerance in oil crops.
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    This review examines the complex regulatory network governing seed oil biosynthesis, including the LAFL transcription factor network, a potential Nuclear Factor-Y regulatory network and the influences of environment factors on seed oil biosynthesis. Additionally, sugar transport, lipid metabolism, fatty acid biosynthesis inhibitors and other fine-tuning mechanisms influence oil content.
      
    Simultaneous mutations in ITPK4 and MRP5 genes result in a low phytic acid level without compromising salt tolerance in Arabidopsis
    Yuying Ren, Mengdan Jiang, Jian-Kang Zhu, Wenkun Zhou, Chunzhao Zhao
    J Integr Plant Biol 2024, 66 (10): 2109-2125.  
    DOI: 10.1111/jipb.13745
    Abstract (Browse 451)  |   Save
    Generation of crops with low phytic acid (myo-inositol-1,2,3,4,5,6-hexakisphosphate (InsP6)) is an important breeding direction, but such plants often display less desirable agronomic traits. In this study, through ethyl methanesulfonate-mediated mutagenesis, we found that inositol 1,3,4-trisphosphate 5/6-kinase 4 (ITPK4), which is essential for producing InsP6, is a critical regulator of salt tolerance in Arabidopsis. Loss of function of ITPK4 gene leads to reduced root elongation under salt stress, which is primarily because of decreased root meristem length and reduced meristematic cell number. The itpk4 mutation also results in increased root hair density and increased accumulation of reactive oxygen species during salt exposure. RNA sequencing assay reveals that several auxin-responsive genes are down-regulated in the itpk4-1 mutant compared to the wild-type. Consistently, the itpk4-1 mutant exhibits a reduced auxin level in the root tip and displays compromised gravity response, indicating that ITPK4 is involved in the regulation of the auxin signaling pathway. Through suppressor screening, it was found that mutation of Multidrug Resistance Protein 5 (MRP5)5 gene, which encodes an ATP-binding cassette (ABC) transporter required for transporting InsP6 from the cytoplasm into the vacuole, fully rescues the salt hypersensitivity of the itpk4-1 mutant, but in the itpk4-1 mrp5 double mutant, InsP6 remains at a very low level. These results imply that InsP6 homeostasis rather than its overall amount is beneficial for stress tolerance in plants. Collectively, this study uncovers a pair of gene mutations that confer low InsP6 content without impacting stress tolerance, which offers a new strategy for creating “low-phytate” crops.
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    Engineering of crops with low phytic acid (InsP6) often impairs other agronomic traits. Simultaneous mutations in the inositol 1,3,4-trisphosphate 5/6-kinase gene ITPK4 and the ABC transporter gene MRP5 result in reduced InsP6 content without affecting plant salt tolerance, providing a promising strategy for breeding crops with low InsP6.
      
    PIF4 interacts with ABI4 to serve as a transcriptional activator complex to promote seed dormancy by enhancing ABA biosynthesis and signaling
    Xiaofeng Luo, Yujia Dai, Baoshan Xian, Jiahui Xu, Ranran Zhang, Muhammad Saad Rehmani, Chuan Zheng, Xiaoting Zhao, Kaitao Mao, Xiaotong Ren, Shaowei Wei, Lei Wang, Juan He, Weiming Tan, Junbo Du, Weiguo Liu, Shu Yuan and Kai Shu
    J Integr Plant Biol 2024, 66 (5): 909-927.  
    doi: 10.1111/jipb.13615
    Abstract (Browse 506)  |   Save
    Transcriptional regulation plays a key role in the control of seed dormancy, and many transcription factors (TFs) have been documented. However, the mechanisms underlying the interactions between different TFs within a transcriptional complex regulating seed dormancy remain largely unknown. Here, we showed that TF PHYTOCHROME-INTERACTING FACTOR4 (PIF4) physically interacted with the abscisic acid (ABA) signaling responsive TF ABSCISIC ACID INSENSITIVE4 (ABI4) to act as a transcriptional complex to promote ABA biosynthesis and signaling, finally deepening primary seed dormancy. Both pif4 and abi4 single mutants exhibited a decreased primary seed dormancy phenotype, with a synergistic effect in the pif4/abi4 double mutant. PIF4 binds to ABI4 to form a heterodimer, and ABI4 stabilizes PIF4 at the protein level, whereas PIF4 does not affect the protein stabilization of ABI4. Subsequently, both TFs independently and synergistically promoted the expression of ABI4 and NCED6, a key gene for ABA anabolism. The genetic evidence is also consistent with the phenotypic, physiological and biochemical analysis results. Altogether, this study revealed a transcriptional regulatory cascade in which the PIF4–ABI4 transcriptional activator complex synergistically enhanced seed dormancy by facilitating ABA biosynthesis and signaling.
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    The transcription factors PIF4 and ABI4 form a complex and synergistically enhance seed dormancy by promoting abscisic acid biosynthesis and signaling. ABI4 stabilizes PIF4 protein, and both transcription factors independently and collaboratively promote ABI4 and NCED6 transcription
      
    Potassium transporter OsHAK9 regulates seed germination under salt stress by preventing gibberellin degradation through mediating OsGA2ox7 in rice
    Peng Zeng, Ting Xie, Jiaxin Shen, Taokai Liang, Lu Yin, Kexin Liu, Ying He, Mingming Chen, Haijuan Tang, Sunlu Chen, Sergey Shabala, Hongsheng Zhang and Jinping Cheng
    J Integr Plant Biol 2024, 66 (4): 731-748.  
    doi: 10.1111/jipb.13642
    Abstract (Browse 444)  |   Save
    Soil salinity has a major impact on rice seed germination, severely limiting rice production. Herein, a rice germination defective mutant under salt stress (gdss) was identified by using chemical mutagenesis. The GDSS gene was detected via MutMap and shown to encode potassium transporter OsHAK9. Phenotypic analysis of complementation and mutant lines demonstrated that OsHAK9 was an essential regulator responsible for seed germination under salt stress. OsHAK9 is highly expressed in germinating seed embryos. Ion contents and non-invasive micro-test technology results showed that OsHAK9 restricted K+ efflux in salt-exposed germinating seeds for the balance of K+/Na+. Disruption of OsHAK9 significantly reduced gibberellin 4 (GA4) levels, and the germination defective phenotype of oshak9a was partly rescued by exogenous GA3 treatment under salt stress. RNA sequencing (RNA-seq) and real-time quantitative polymerase chain reaction analysis demonstrated that the disruption of OsHAK9 improved the GA-deactivated gene OsGA2ox7 expression in germinating seeds under salt stress, and the expression of OsGA2ox7 was significantly inhibited by salt stress. Null mutants of OsGA2ox7 created using clustered, regularly interspaced, short palindromic repeat (CRISPR)/CRISPR-associated nuclease 9 approach displayed a dramatically increased seed germination ability under salt stress. Overall, our results highlight that OsHAK9 regulates seed germination performance under salt stress involving preventing GA degradation by mediating OsGA2ox7, which provides a novel clue about the relationship between GA and OsHAKs in rice.
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    The causal gene of the rice (Oryza sativa) germination defective under salt stress mutant encodes the potassium transporter OsHAK9, which reduces K+ loss, promotes Na+ accumulation, and inhibits OsGA2ox7 expression to prevent gibberellin degradation, thus significantly enhancing seed germination under salt stress.
      
    Orchestrating seed storage protein and starch accumulation toward overcoming yield–quality trade-off in cereal crops
    Shuanghe Cao, Bingyan Liu, Daowen Wang, Awais Rasheed, Lina Xie, Xianchun Xia and Zhonghu He
    J Integr Plant Biol 2024, 66 (3): 468-483.  
    DOI: 10.1111/jipb.13633
    Abstract (Browse 461)  |   Save
    Achieving high yield and good quality in crops is essential for human food security and health. However, there is usually disharmony between yield and quality. Seed storage protein (SSP) and starch, the predominant components in cereal grains, determine yield and quality, and their coupled synthesis causes a yield–quality trade-off. Therefore, dissection of the underlying regulatory mechanism facilitates simultaneous improvement of yield and quality. Here, we summarize current findings about the synergistic molecular machinery underpinning SSP and starch synthesis in the leading staple cereal crops, including maize, rice and wheat. We further evaluate the functional conservation and differentiation of key regulators and specify feasible research approaches to identify additional regulators and expand insights. We also present major strategies to leverage resultant information for simultaneous improvement of yield and quality by molecular breeding. Finally, future perspectives on major challenges are proposed.
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    This review compares the molecular machinery orchestrating seed storage protein and starch accumulation in maize, rice, and wheat; proposes approaches to identify additional regulators coupling seed storage protein and starch biosynthesis; and provides perspectives on key challenges for synchronous improvement of cereal yield and quality.
      
    Antagonistic MADS-box transcription factors SEEDSTICK and SEPALLATA3 form a transcriptional regulatory network that regulates seed oil accumulation
    Shuangcheng He, Yuanchang Min, Zijin Liu, Fang Zhi, Rong Ma, Ankang Ge, Shixiang Wang, Yu Zhao, Danshuai Peng, Da Zhang, Minshan Jin, Bo Song, Jianjun Wang, Yuan Guo and Mingxun Chen
    J Integr Plant Biol 2024, 66 (1): 121-142.  
    DOI: 10.1111/jipb.13606
    Abstract (Browse 517)  |   Save
    Transcriptional regulation is essential for balancing multiple metabolic pathways that influence oil accumulation in seeds. Thus far, the transcriptional regulatory mechanisms that govern seed oil accumulation remain largely unknown. Here, we identified the transcriptional regulatory network composed of MADS-box transcription factors SEEDSTICK (STK) and SEPALLATA3 (SEP3), which bridges several key genes to regulate oil accumulation in seeds. We found that STK, highly expressed in the developing embryo, positively regulates seed oil accumulation in Arabidopsis (Arabidopsis thaliana). Furthermore, we discovered that SEP3 physically interacts with STK in vivo and in vitro. Seed oil content is increased by the SEP3 mutation, while it is decreased by SEP3 overexpression. The chromatin immunoprecipitation, electrophoretic mobility shift assay, and transient dual-luciferase reporter assays showed that STK positively regulates seed oil accumulation by directly repressing the expression of MYB5, SEP3, and SEED FATTY ACID REDUCER 4 (SFAR4). Moreover, genetic and molecular analyses demonstrated that STK and SEP3 antagonistically regulate seed oil production and that SEP3 weakens the binding ability of STK to MYB5, SEP3, and SFAR4. Additionally, we demonstrated that TRANSPARENT TESTA 8 (TT8) and ACYL-ACYL CARRIER PROTEIN DESATURASE 3 (AAD3) are direct targets of MYB5 during seed oil accumulation in Arabidopsis. Together, our findings provide the transcriptional regulatory network antagonistically orchestrated by STK and SEP3, which fine tunes oil accumulation in seeds.
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    SEEDSTICK and SEPALLATA3 antagonistically regulate seed oil accumulation in Arabidopsis; this lays the foundation for engineering high-oil cultivars of oil-producing crops.
      
    TaSRO1 interacts with TaVP1 to modulate seed dormancy and pre-harvest sprouting resistance in wheat
    Shupeng Liu, Li Li, Wenlong Wang, Guangmin Xia and Shuwei Liu
    J Integr Plant Biol 2024, 66 (1): 36-53.  
    DOI: 10.1111/jipb.13600
    Abstract (Browse 443)  |   Save
    Dormancy is an adaptive trait which prevents seeds from germinating under unfavorable environmental conditions. Seeds with weak dormancy undergo pre-harvest sprouting (PHS) which decreases grain yield and quality. Understanding the genetic mechanisms that regulate seed dormancy and resistance to PHS is crucial for ensuring global food security. In this study, we illustrated the function and molecular mechanism of TaSRO1 in the regulation of seed dormancy and PHS resistance by suppressing TaVP1. The tasro1 mutants exhibited strong seed dormancy and enhanced resistance to PHS, whereas the mutants of tavp1 displayed weak dormancy. Genetic evidence has shown that TaVP1 is epistatic to TaSRO1. Biochemical evidence has shown that TaSRO1 interacts with TaVP1 and represses the transcriptional activation of the PHS resistance genes TaPHS1 and TaSdr. Furthermore, TaSRO1 undermines the synergistic activation of TaVP1 and TaABI5 in PHS resistance genes. Finally, we highlight the great potential of tasro1 alleles for breeding elite wheat cultivars that are resistant to PHS.
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    In wheat (Triticum aestivum), the small protein SIMILAR TO RCD ONE (TaSRO1) regulates seed dormancy and resistance to pre-harvest sprouting by interacting with and suppressing the function of TaViviparous1.
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