Scientia Agricultura Sinica ›› 2020, Vol. 53 ›› Issue (5): 857-873.doi: 10.3864/j.issn.0578-1752.2020.05.001

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

ABA Metabolism and Signaling and Their Molecular Mechanism Regulating Seed Dormancy and Germination

SONG SongQuan1,4,LIU Jun2,XU HengHeng2,LIU Xu3(),HUANG Hui4   

  1. 1 Institute of Botany, Chinese Academy of Sciences, Beijing 100093
    2 Agro-Biological Gene Research Center, Guangdong Academy of Agricultural Sciences, Guangzhou 510640
    3 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081
    4 Key Laboratory of Research and Utilization of Ethnomedicinal Plant Resources of Hunan Province, Huaihua University/College of Biological and Food Engineering, Huaihua 418008, Hunan
  • Received:2019-05-21 Accepted:2019-09-30 Online:2020-03-01 Published:2020-03-14
  • Contact: Xu LIU E-mail:liuxu01@caas.cn

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Abstract:

Seed dormancy is an adaptive characteristic to environmental changes acquired by many plants during long-term phylogenetic development, and is an effective way regulating the optimal spatiotemporal distribution of seed germination and seedling formation, and is also a selective strategy for the successful reproduction and propagation in species. Phytohormonal regulation of seed dormancy and germination may be a highly conserved mechanism, of which abscisic acid (ABA) plays a master role in dormancy release and germination, and gibberellin (GA) functions as stimulating seed germination after dormancy is released. The role of ABA in seed dormancy and germination is mainly regulated by its metabolism (biosynthesis and catabolism) and signaling pathways. Therefore, in this paper, we mainly summarize the research progresses of ABA metabolism and signaling, the effects of ABA on seed development, dormancy and germination as well as the relationships between DOG1 (DELAY OF GERMINATION1, a specific gene involved in seed dormancy) and ABA signaling components. The researches showed that C40 epoxycarotenoid is a precursor, and zeaxanthin epoxidase and 9-cis-epoxycarotenoid dioxygenase are the principal regulatory enzymes in ABA biosynthesis. The ABA catabolism includes hydroxylation and conjugation with glucose. The hydroxylation of ABA at C-8' position is catalyzed by the CYP707A, which is an important step for ABA catabolism. In the core ABA signaling pathway, ABA binds to PYR/PYL/RCAR receptors and triggers a conformational change that allows receptor-ABA complex to bind to and inhibit type 2C protein phosphatase (PP2C) activity, which results in de-repression and activation of kinases such as sucrose non-fermenting1-related protein kinase 2 (SnRK2). These kinases then phosphorylate and activate transcription factors (TF), which bind to the target promoters and induce the expression of ABA response gene downstream. ABA accumulates in seeds during mid- and late-maturation stages, and ABA synthesized in zygotic tissues induces primary dormancy and promotes seed maturation. ABA content accumulated during development and preserved in dry seeds declines at the early stage of seed imbibition. ABA is a positive regulator of seed dormancy induction and maintenance, and is a negative regulator of seed germination. DOG1 expresses and functions during seed maturation, and its expression is regulated by alternative splicing and alternative polyadenylation. Antisense DOG1 is a repressor of seed dormancy, which negatively regulates DOG1 expression and seed dormancy by causing transcriptional interference and affecting transcription extension. Seed dormancy and germination are regulated not only by core ABA signaling pathway, but also by DOG1-AHG1 (ABA HYPERSENSITIVE GERMINATION1)/AHG3 pathway. DOG1 can bind to AHG1/AHG3 and cause seed dormancy by sequestrating those negative regulators of ABA signaling and increasing ABA sensitivity in seeds. Finally, we propose some scientific issues required for investigation further in the future. How do ABA 8'-hydroxylase, ABA glucosyltransferase and β-glucosidase and their genes respond to developmental and environmental changes to maintain the normal ABA levels in ABA catabolism? How do the important regulators in ABA physiology such as Ca 2+ or reactive oxygen species influence the core ABA signaling pathway? Which pathway is preferentially responded by PP2C, a downstream overlapping component of core ABA signaling pathway and DOG1-AHG1/AHG3 pathway, when it integrates physiological conditions or environmental signals, and how are these two pathways coordinated, and what new target components does PP2C have? This paper will provide a basis to further investigate the molecular mechanism regulating seed dormancy and germination by ABA.

Key words: abscisic acid, dormancy, dormancy gene DOG1, germination, metabolism, signaling

Fig. 1

ABA biosynthetic and catabolic pathways (Modified from DEJONGHE et al.[13]) ABA precursor is synthesized from the methylerythritol phosphate (MEP) pathway. Enzymes are shown in red colour. ZEP: Zeaxanthin epoxidase; NSY: Neoxanthin synthase; NCED: 9-cis-epoxycarotenoid dioxygenase; XD: Xanthoxin dehydrogenase; ABAO: Abscisic aldehyde oxidase; CYP707A: ABA 8'-hydroxylase; ABH1: Phaseic acid reductase 1; ABAGT: ABA glucosyltransferase; βG: β-glucosidase. Enzyme inhibitors are shown in blue colour. (+)-9'-AABA: (+)-9'-acetylene-ABA; AHI4: ABA-8'-hydroxylase inhibitor 4; (+)-8'-MABA: (+)-8'-methylidyne-ABA; NDGA: Nordihydroguaiaretic acid; SLCCD13: Sesquiterpene-like carotenoid cleavage dioxygenase inhibitor 13"

Fig. 2

ABA-induced changes in receptor conformation (From CUTLER et al.[12]) In the absence of ABA, PYR/PYL (pyrabactin resistance 1/pyrabactin resistance 1-like) proteins possess an open conformation of the gate and latch loops (red and green, respectively) that flank the ABA-binding pocket. Binding of ABA induces closure of the gate and latch, which in turn creates the interaction surface that recruits docking of type 2C protein phosphatases (PP2C) onto the ABA-bound receptors. A conserved proline in the gate (which corresponds to the residue to proline 88 in PYR1 and is shown in blue) forms a direct contact with the PP2C at the docking site, which explains the PP2C-binding defect observed with PYR1P88"

Fig. 3

ABA signalling pathway and emerging model of seed dormancy regulated by DOG1 (Modified from NONOGAKI [19]) In the ABA perception and signaling pathway (left), ABA receptors (PYR/PYL/RCARs) bind to and inactivate the ABA INSENSITIVE1 (ABI1) subfamily protein phosphatases 2C (PP2Cs), including ABI1, ABI2, HYPERSENSITIVE TO ABA1 (HAB1) and HAB2, which results in de-repression and activation of kinases, such as sucrose nonfermenting1-related protein kinase 2 (SnRK2). These kinases then phosphorylate and activate transcription factors (TF), which bind to the target promoters (Pro), to induce ABA-responsive genes downstream. For seed dormancy regulation (right), DOG1 binds to ABA HYPERSENSITIVE GERMINATION1 (AHG1) and AHG3, PP2Cs primarily functioning in seeds. The DOG1 is thought to cause seed dormancy by sequestrating these negative regulators of ABA signaling and increasing ABA sensitivity in seeds"

Fig. 4

Regulation of DOG1 expression and function (From NONOGAKI[91]) A: Structures of the DOG1 gene. Top: DOG1 gDNA with exons (E1, E2, E3) and introns (I1, I2). Alternatively spliced regions are highlighted in pink and orange. Approximate positions of the dog1 mutations (T-DNAs in dog1-3, dog1-4, dog1-5 and a single-base deletion [-C] in dog1-1) are also indicated. Middle: Alternative DOG1 transcripts (α, β, γ, δ, ε) and the corresponding proteins. Note that DOG1-ε is not exactly an alternative splicing product. Bottom: Alternatively polyadenylated short DOG1 (shDOG1), which is identical to DOG1-ε and long (lgDOG1) transcripts, which comprises DOG1-α, -β, -γ and -δ. The transcriptional start (TSS) and termination (TTS) sites are indicated. Approximate position and the orientation of antisense DOG1 (asDOG1) are shown as a blue arrow. B: Possible mechanisms of asDOG1 function. Relatively stable asDOG1 RNA could function as a regulatory RNA, in a sequence-specific manner or through its secondary structure, for RNA-mediated chromatin remodeling (right panel, trans regulation). However, allele-specific asDOG1 expression has indicated that asDOG1 functions in cis (left panel). The “act” of transcription itself, rather than its product (RNA), exerts the negative effects of asDOG1 expression to DOG1 expression and dormancy. Antisense expression could cause transcriptional interference and affect transcription elongation, which is known to be important for DOG1 expression and seed dormancy while transcription-mediated chromatin remodeling is also possible. AS: Alternative splicing; APA: Alternative polyadenylation; Dist: Distal; Prox: Proximal; Prot: Protein; Tran: Transcription"

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