脱落酸调控种子休眠和萌发的分子机制

doi:10.11983/CBB18080

摘要/Abstract

摘要： 脱落酸(ABA)是调控种子休眠和萌发过程的主要植物激素。种子内源ABA含量和种胚对ABA敏感性共同调控种子休眠和萌发过程, 确保植物种子以休眠状态在逆境中保持其自身繁衍能力, 并在适宜的环境下启动萌发程序。种子ABA合成代谢和ABA信号转导途径涉及许多重要基因家族, 它们通过复杂的调控网络精确地控制着种胚发生、种子成熟、休眠及萌发进程。该文对ABA调控种子休眠和萌发的分子机制最新研究进展进行综述, 并展望了今后的研究方向。

关键词: 脱落酸, 种子休眠, 种子萌发

Abstract: Abscisic acid (ABA) acts as a major phytohormone in regulating seed dormancy and germination. Endogenous ABA level in the seed and embryo sensitivity to ABA jointly regulate the seed dormancy and germination processes, ensuring that the dormant seed can maintain self-reproductive ability under adverse situations and germinate under optimal conditions. A number of key genes are involved in regulating the ABA metabolism and signaling transduction pathway, and these genes accurately control the processes of embryogenesis, seed maturation, dormancy, and germination via a complicated regulation network. In this paper, we review the latest research in the molecular mechanism of ABA-regulated seed dormancy and germination and pinpoint prospects for future research.

Key words: abscisic acid, seed dormancy, seed germination

伍静辉, 谢楚萍, 田长恩, 周玉萍. 脱落酸调控种子休眠和萌发的分子机制. 植物学报, 2018, 53(4): 542-555.

Wu Jinghui, Xie Chuping, Tian Changen, Zhou Yuping. Molecular Mechanism of Abscisic Acid Regulation During Seed Dormancy and Germination. Chinese Bulletin of Botany, 2018, 53(4): 542-555.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB18080

https://www.chinbullbotany.com/CN/Y2018/V53/I4/542

图/表 2

表1 参与种子休眠的关键基因

Table 1 Key genes involved in seed dormancy

基因名称	突变体休眠能力	基本功能	参考文献
NCED6/9	变化不显著	ABA合成途径关键代谢酶	Lefebvre et al., 2006
CYP707A1/2	增强	ABA代谢途径关键代谢酶, 其表达受ABI4调控	Kushiro et al., 2004; Okamoto et al., 2006; Shu et al., 2013
ABI3	降低	促进种子发育, 抑制种子萌发	Raz et al., 2001; Dekkers et al., 2016
FUS3	降低	促进种子发育, 增强NCED6和NCED9的表达	Raz et al., 2001; Tiedemann et al., 2008; Yamamoto et al., 2010
LEC2	降低	促进种子发育, 促进LEC1、FUC3和DOG1的表达	Raz et al., 2001; Braybrook et al., 2006; Stone et al., 2008
LEC1/L1L	降低	促进种子发育, 激活ABI3、FUS3和LEC2的表达	Raz et al., 2001; Kagaya et al., 2005; To et al., 2006
DEP	降低	促进ABI3的表达	Barrero et al., 2010
DOG1	降低	种子休眠正向调控因子	Née et al., 2017
ABI4	降低	促进种子发育, 是ABA/GA的关键调控因子	Shu et al., 2013
SPT	Col背景中增强; Ler背景中降低	调控ABI4表达, 在Col和Ler背景下功能相反	Belmonte et al., 2013; Vaistij et al., 2013
CHO1	降低	促进ABA合成, 抑制ABA降解	Yano et al., 2009
MYB96	降低	促进种子休眠, 抑制种子萌发	Lee et al., 2015a, 2015b
ABI5	变化不显著	不直接参与种子休眠调控, 但负调控种子萌发	Piskurewicz et al., 2008; Kanai et al., 2010
bZIP10/25/53	降低	协同促进种子发育	Alonso et al., 2009
WRKY41	降低	促进ABI3转录	Ding et al., 2014
DAG1	降低	促进ABA而抑制GA积累, 正调控种子休眠	Boccaccini et al., 2014, 2016
DOF6	未提及	与RGL2构成复合体, 共同促进种子初级休眠	Ravindran et al., 2017
KYP/SUVH4	增强	抑制DOG1和ABI3转录, 负调控种子休眠	Zheng et al., 2012
LDL1/LDL2	增强	抑制DOG1表达, 负调控种子休眠	Zhao et al., 2015

表1 参与种子休眠的关键基因

Table 1 Key genes involved in seed dormancy

基因名称	突变体休眠能力	基本功能	参考文献
NCED6/9	变化不显著	ABA合成途径关键代谢酶	Lefebvre et al., 2006
CYP707A1/2	增强	ABA代谢途径关键代谢酶, 其表达受ABI4调控	Kushiro et al., 2004; Okamoto et al., 2006; Shu et al., 2013
ABI3	降低	促进种子发育, 抑制种子萌发	Raz et al., 2001; Dekkers et al., 2016
FUS3	降低	促进种子发育, 增强NCED6和NCED9的表达	Raz et al., 2001; Tiedemann et al., 2008; Yamamoto et al., 2010
LEC2	降低	促进种子发育, 促进LEC1、FUC3和DOG1的表达	Raz et al., 2001; Braybrook et al., 2006; Stone et al., 2008
LEC1/L1L	降低	促进种子发育, 激活ABI3、FUS3和LEC2的表达	Raz et al., 2001; Kagaya et al., 2005; To et al., 2006
DEP	降低	促进ABI3的表达	Barrero et al., 2010
DOG1	降低	种子休眠正向调控因子	Née et al., 2017
ABI4	降低	促进种子发育, 是ABA/GA的关键调控因子	Shu et al., 2013
SPT	Col背景中增强; Ler背景中降低	调控ABI4表达, 在Col和Ler背景下功能相反	Belmonte et al., 2013; Vaistij et al., 2013
CHO1	降低	促进ABA合成, 抑制ABA降解	Yano et al., 2009
MYB96	降低	促进种子休眠, 抑制种子萌发	Lee et al., 2015a, 2015b
ABI5	变化不显著	不直接参与种子休眠调控, 但负调控种子萌发	Piskurewicz et al., 2008; Kanai et al., 2010
bZIP10/25/53	降低	协同促进种子发育	Alonso et al., 2009
WRKY41	降低	促进ABI3转录	Ding et al., 2014
DAG1	降低	促进ABA而抑制GA积累, 正调控种子休眠	Boccaccini et al., 2014, 2016
DOF6	未提及	与RGL2构成复合体, 共同促进种子初级休眠	Ravindran et al., 2017
KYP/SUVH4	增强	抑制DOG1和ABI3转录, 负调控种子休眠	Zheng et al., 2012
LDL1/LDL2	增强	抑制DOG1表达, 负调控种子休眠	Zhao et al., 2015

图1 以PYR/PYL/RCAR和ABAR/CHLH为受体的ABA信号转导通路模型细胞质ABA受体PYR/PYL/RCAR在静息状态下以二聚体形式存在, 当与ABA结合后则以单体形式与PP2Cs结合, 解除对SnRK2s的抑制, 磷酸化ABI5及RAV1等转录因子, 激活下游ABA响应基因。ABAR/CHLH是叶绿体膜上的ABA受体, 其C-端和N-端暴露在细胞质中。低浓度ABA时, 叶绿体内伴侣蛋白CPN20结合ABAR, 抑制ABAR与负调控因子WRKY40的相互作用, 同时WRKY40与WRKY18和WRKY60通过直接或间接作用, 抑制ABF4/ABI4/ABI5的表达。高浓度ABA促使WRKY40向细胞质迁移, ABAR的C-端与WRKY40相互作用, 解除WRKY40对ABF4/ABI4/ABI5的转录抑制, 实现ABA生理效应。RAV1通过抑制ABI3/ABI4/ABI5转录来负调控ABA信号; WRKY6、WRKY41和NF-YC-RGL2分别通过下调RAV1的表达, 促进ABI3和ABI5的表达, 它们均正调控ABA信号。图中(+)和(-)分别表示该基因在ABA信号通路中具有正向和反向调控作用。

Figure 1 Models of ABA signaling transduction pathway based on ABA receptors PYR/PYL/RCAR and ABAR/CHLHIntracellular ABA receptor PYR/PYL/RCAR exists as a homodimer in the resting state. Upon binding to ABA, PYR/PYL/RCAR turns into monomer and interacts with PP2Cs to release SnRK2s, which phosphorylates transcription factors like ABI5 and RAV1, to activate downstream ABA responsive genes. ABAR/CHLH is a protein across the chloroplast membrane, whose C- and N- ends are exposed to the cytoplasm. In the absence or low levels of ABA, the interaction of CPN20 with ABAR attenuates the interaction between ABAR and the negative regulator WRKY40, meanwhile WRKY40, WRKY18 and WRKY60 directly or indirectly inhibit the expression of ABF4/ABI4/ABI5. In response to a high level of ABA, WRKY40 is recruited from nucleus to the cytosol. C-terminal part of ABA-bound ABAR interacts with WRKY40 and relieves the suppression of ABF4/ABI4/ABI5 by WRKY40, promoting the physiological effects of ABA. RAV1 negatively regulates the ABA signaling by inhibiting the transcription of ABI3/ABI4/ABI5. WRKY6, WRKY41, and NF-YC-RGL2 play the positive roles in the ABA signaling, by down-regulating the expression of RAV1, promoting the expression of ABI3 and ABI5, respectively. (+) and (-) in this figure denote the positive regulation and the negative regulation in the ABA signaling pathway, respectively.

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