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Acta Agronomica Sinica ›› 2024, Vol. 50 ›› Issue (1): 1-15.doi: 10.3724/SP.J.1006.2024.34012

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Research progress of seed dormancy and germination regulation

SONG Song-Quan1,3,*(), TANG Cui-Fang2,3, LEI Hua-Ping3, JIANG Xiao-Cheng2, WANG Wei-Qing1, CHENG Hong-Yan1   

  1. 1Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
    2College of Life Sciences, Hunan Normal University, Changsha 410081, Hunan, China
    3Nanling Research Institute for Modern Seed Industry, Xiangnan University, Chenzhou 423099, Hunan, China
  • Received:2022-12-18 Accepted:2023-09-08 Online:2024-01-12 Published:2023-09-08
  • Contact: *E-mail: sqsong2019@163.com
  • Supported by:
    National Key Technology Support Program of China(2012BAC01B05);Provincial Special Project of Chenzhou National Sustainable Development Agenda Innovation Demonstration Zone Construction(2022sfq06);Chenzhou Municipal People’s Government and Xiangnan University Talent Project

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Abstract:

Dormancy enables plant seeds to time germination until environmental conditions become favorable for seedling survival and growth. The dormancy characteristics of seeds are of important ecological adaptive significance and notable agricultural value. Phytohormone abscisic acid (ABA) and gibberellin (GA) are the key factors for seed dormancy and germination. Mature seeds in dormancy state contain high levels of ABA and low levels of GA. ABA induces and maintains seed dormancy, while GA antagonizes ABA and promotes seed germination. DELAY OF GERMINATION-1 (DOG1) is a major regulator of seed dormancy and had a synergistic effect with ABA to delay germination. DOG1 enhances ABA signal transduction by combining with PP2C ABA hypersensitive germination (AHG1/AHG3), and inhibits the action of AHG1 to increase ABA sensitivity and impose seed dormancy. Imprinted genes are regulated by epigenetic mechanisms before and after fertilization, and are closely related to the establishment and release of seed dormancy. In recent years, remarkable progress has been made in the regulation of seed dormancy. In the present paper, we reviewed the effects of phytohormones ABA and GA on seed dormancy and germination, the action mechanism regulating seed dormancy by DOG1, and the epigenetic regulation of seed dormancy and germination. In addition, we also propose some scientific issues that need to be further investigated in this field to provide some information for understanding the molecular mechanism of seed dormancy and germination, breeding in anti-preharvest sprouting in crop plants, and promoting the germination of dormant seeds.

Key words: DOG1, epigenetic, molecular mechanism, phytohormone, seed dormancy

Fig. 1

Regulatory phytohormone networks in seed dormancy and germination Three major phytohormones, including abscisic acid (ABA), gibberellin (GA), and auxin, are key players in seed dormancy and germination. Mature seeds are dormant and contain a high level of ABA and a low level of GA. Several transcription factors (ABI4, DDF1, OsAP2-39, AP2, and CHO1) are involved at seed dormancy stage by positively regulating (+) the accumulation of ABA and decreasing GA content. While seed dormancy is broken, the seed becomes non-dormant and the initiation of germination can start. At this stage, the ABA/GA balance is kept by positive and negative regulation signals of almost all other phytohormones, including ethylene (ET), brassinosteroid (BR), jasmonic acid (JA), salicylic acid (SA), cytokinins (CTK), and strigolactones (SL). Transcription factors, including ARF, MYB96, ABI3, ABI4, and ABI5, regulate ABA biosynthesis by interacting with CYP707A1 and CYP707A2, while GA-negative regulation (-) is ensured by DELLA genes. The balance is constantly maintained until the seed emergence step. These information are from Sohn et al.[8]"

Fig. 2

A model for the transcriptional regulation of DOG1 gene (promotor in red; coding region in green) to induce dormancy in Arabidopsis thaliana seeds Downstream of the GBL1 cis-element is INDEL (insertion-deletion), which is a 285 bp sequence. INDEL is present in DOG1 promoter of Ler-0 accession (weakly dormant). However, INDEL is absent in Cvi-0 ecotype (strong dormant). INDEL directly affects the ability of bZIP67 to trans-activate DOG1 in vivo and may explain the observed variation in DOG1 transcript levels and consequently dormancy between ecotypes. Such natural allelic variation in DOG1 coupled with DOG1 expression plasticity confers substantial adaptive significance in the field, where seasonal environmental factors can vary greatly, and the optimal timing of seed germination is primordial. ETR1: ethylene response-1; DRE/CRT: dehydration-responsive element/C-repeat; TPL1: topless-1; ERF12: ethylene responsive transcription factor-12; GBL1 and GBL2: G-box-like-1 and -2; SOM: somnus. These information are from Carrillo-Barral et al. [17]"

Fig. 3

Model depicting how different combinations of epigenetic modifications on parental alleles affect gene expression in the endosperm and control seed dormancy Genes whose maternal alleles are suppressed by FIS-PRC2-mediated H3K27me3 encode ethylene-pathway genes that are necessary to induce germination. The paternal alleles of those genes are likely suppressed by CHHm established by the non-canonical RdDM pathway. Cold stress-inducible AGO6 may also target maternal alleles and apply cold stress-responsive CHHm by the non-canonical RdDM pathway. REF6 binds to the CTCTGYTY motif (Y = C or T) to activate the target genes. In dormant seeds, the maternal alleles are activated by the H3K27me3 demethylase REF6 and the genes show maternally biased expression patterns. In non-dormant seeds, the genes are biallelically expressed, likely because the RdDM pathway is specifically active under dormancy-inducing conditions. FIS-PRC2, SUVH family proteins, and CMT3 establish H3K27me3, H3K9me2, and CHGm, respectively, on maternal alleles of PEGs. Because CHGm on the REF6-binding site inhibits binding activity of REF6, the maternal alleles are continuously silenced, and the genes reveal paternally biased gene expression patterns throughout endosperm development including germination. ABI3, an important transcription factor establishing seed dormancy belongs to this category. Genes with single H3K9me2 on the maternal alleles are fully silenced throughout development. These maternal-specific epigenetic modifications are detected in developing endosperm at 4 days after pollination. These information are from Sato and K?hler[2]."

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