重要的种子储存物质长寿命mRNA

doi:10.11983/CBB24006

植物学报 ›› 2024, Vol. 59 ›› Issue (3): 355-372.DOI: 10.11983/CBB24006

重要的种子储存物质长寿命mRNA

朱晓博¹^,², 董张¹^,², 祝梦瑾¹^,², 胡晋¹^,², 林程³, 陈敏², 关亚静¹^,²^,^*()

¹浙江大学海南研究院, 三亚 572000
²浙江大学农业与生物技术学院现代种业研究所, 杭州 310058
³苏州农业职业技术学院, 苏州 215008

收稿日期:2024-01-10 接受日期:2024-03-30 出版日期:2024-05-01 发布日期:2024-05-08
通讯作者: *关亚静, 博士, 教授、博导, 浙江大学“求是”青年学者, 农业与生物技术学院现代种业研究所种子科学与工程中心主任, 浙江大学海南研究院种子工程与检验检疫团队方向负责人, 中国作物学会种子专业委员会副会长, 中国植物学会种子与技术专业委员会副主任。主要从事种子发育成熟和活力分子机理、多功能种子丸化及种子增值技术研究。E-mail: vcguan@zju.edu.cn
基金资助:
海南省科技计划三亚崖州科技城联合项目(320LH032);海南省自然科学基金(322CXTD522);浙江省“三农九方”科技协作计划(2023SNJF04302);苏州市农业科技创新项目(SNG2022063);江苏省“双创博士“项目(JSSCBS20221012)

Indispensable Material for Germination: Long-lived mRNAs of Plant Seed

Xiaobo Zhu¹^,², Zhang Dong¹^,², Mengjin Zhu¹^,², Jin Hu¹^,², Cheng Lin³, Min Chen², Yajing Guan¹^,²^,^*()

¹Hainan Institute, Zhejiang University, Sanya 572000, China
²The Advanced Seed Institute, College of Agriculture and Biotechnology, Zhejiang University, Hangzhou 310058, China
³Suzhou Polytechnic Institute of Agriculture, Suzhou 215008, China

Received:2024-01-10 Accepted:2024-03-30 Online:2024-05-01 Published:2024-05-08
Contact: *E-mail: vcguan@zju.edu.cn

1. 附图1 Act D处理下水稻日本晴种子萌发情况.pdf(221KB)

摘要/Abstract

摘要： 高等植物通常从种子萌发开始, 经过营养生长和生殖发育后重新形成种子, 由此完成世代更迭。种子中积累的碳水化合物、脂质、蛋白质及mRNA等大分子物质对于维持其发芽潜力至关重要, 其中部分mRNA可长期保存而不被降解, 被称为长寿命mRNA (即long-lived mRNA)。在水稻(Oryza sativa)中, 与萌发相关的long-lived mRNA在花后10-20天开始转录积累, 花后20天至种子完全成熟期间, 一些与休眠和胁迫响应相关的long-lived mRNA转录并保存在细胞中。Long-lived mRNA种类繁多, 主要包括蛋白质合成类mRNA、能量代谢类mRNA、细胞骨架类mRNA及逆境响应相关的mRNA, 如小热激蛋白和LEA家族蛋白。Long-lived mRNA的转录组分析表明, 很多基因的启动子区域都包含脱落酸(ABA)或赤霉素(GA)相关的顺式作用元件, 拟南芥(Arabidopsis thaliana) atabi5突变体种子中约有500个不同于野生型的差异表达long-lived mRNA, 暗示ABA和GA是影响long-lived mRNA种类的关键激素。Long-lived mRNA通常与单核糖体和RBP蛋白交联在一起, 以PBs (P-bodies)形式存在于细胞中, 保护mRNA不被降解。与种子休眠相关的long-lived mRNA在种子后熟过程中逐渐被降解, 而且一些特定long-lived mRNA的氧化修饰是种子打破休眠的一种生物现象。在种子长期贮藏过程中, long-lived mRNA的随机降解直接关系到种子的寿命和活力, 保留下来的mRNA在种子吸胀初期被翻译成蛋白质, 促进种子在吸胀早期快速萌发。该文综述了种子重要储存物质long-lived mRNA的特征和功能, 并提出了本领域需要进一步研究的科学问题, 以期为深入理解种子休眠、萌发与寿命的分子机制提供参考。

关键词: 长寿命mRNA, 种子休眠, 种子萌发, 种子贮藏

Abstract: Higher plants usually start from seed germination and re-form seeds after vegetative growth and reproductive development, thus completing the life cycle. Carbohydrates, lipids, proteins, mRNA and other macromolecular substances accumulated in seeds are crucial to maintain the germination potential of seeds, some of mRNA can be preserved for a long time without degradation, known as long-lived mRNA. In rice, long-lived mRNA associated with germination began to be transcribed and accumulated 10 to 20 days after flowering, and some long-lived mRNA associated with dormancy and stress response were transcribed and preserved in cells from 20 days after flowering to seed maturity. There are many kinds of long-lived mRNA, mainly including some protein synthesis mRNA, energy metabolism mRNA, cytoskeleton mRNA and some stress response related mRNA, such as small heat shock protein, LEA (late embryogenesis abundant) family proteins. Transcriptomic analysis found that the promoter regions of many genes contain ABA- or GA-associated cis-acting elements, and there are about 500 differentially expressed long-lived mRNAs in the Arabidopsis atabi5 (ABA-insensitive 5) mutant seeds that differ from the wild type, suggesting that abscisic acid (ABA) and gibberellin (GA) are the key hormones that influence the type of long-lived mRNA. Long-lived mRNAs are usually cross-linked with a single ribosome, RNA binding protein, which exists in cells in the form of P-bodies (PBs) to protect the mRNA from degradation. However, long-lived mRNAs associated with seed dormancy are gradually degraded during seed post-ripening, and the oxidative modification of some specific long-lived mRNAs is also a biological phenomenon to break seed dormancy. During the long-term storage of seeds, the random degradation of long-lived mRNA is directly related to the life and vitality of seeds, and the retained mRNA is translated into protein to help the rapid germination of seeds in the early stage of imbibition. In this paper, the characteristics and functions of long-lived mRNA are reviewed, and some future scientific issues are discussed to provide a reference for further understanding of the molecular mechanisms of seed dormancy, germination and longevity.

Key words: long-lived mRNAs, seed dormancy, seed germination, seed storage

朱晓博, 董张, 祝梦瑾, 胡晋, 林程, 陈敏, 关亚静. 重要的种子储存物质长寿命mRNA. 植物学报, 2024, 59(3): 355-372.

Xiaobo Zhu, Zhang Dong, Mengjin Zhu, Jin Hu, Cheng Lin, Min Chen, Yajing Guan. Indispensable Material for Germination: Long-lived mRNAs of Plant Seed. Chinese Bulletin of Botany, 2024, 59(3): 355-372.

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https://www.chinbullbotany.com/CN/Y2024/V59/I3/355

图/表 3

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基因名称	ID			表达特点		功能	分子机制	物种		参考文献
SDR3.1	LOC_Os03g-11550			开花后5天开始表达, 15天达到峰值		反向调控种子休眠	与ABI5互作, 并抑制ABI5的表达	水稻		Guo et al., 2024
SRO1	TraesCS4A-01G321300			在叶片、根和种子中表达		反向调控种子休眠	与TaVP1互作, 抑制穗上萌发基因TaPHS1和TaSdr的表达, 且影响TaVP1与TaABI5的互作	小麦		Liu et al., 2024
OsWSD1	LOC_Os03g-24460			在种子发育过程中, 其表达量逐渐降低; 在种子吸胀前36小时稳定表达		正向调控种子休眠	抑制GA的合成和α-淀粉酶的活性; 突变后对ABA不敏感	水稻		Huang et al., 2023
AtAIL6	AT5G10510			在整个种子发育过程中维持高表达		正向调控种子休眠	FUS3直接调控AtAIL6的表达	拟南芥		Liu et al., 2023
WRKY36	AT1G69810			在种子中高表达		反向调控种子休眠	WRKY36与AFP2互作, 复合体直接抑制DOG1的表达	拟南芥		Deng et al., 2023
AFP2	AT1G13740			在种子中高表达		反向调控种子休眠	WRKY36与AFP2互作, 复合体直接抑制DOG1的表达	拟南芥		Deng et al., 2023
OsNAC2	LOC_Os04g-38720			在种子发育和萌发过程中高表达		正向调控种子休眠	直接抑制OsABAox1和OsABAox2的表达	水稻		Zhao et al., 2023
BG14	AT2G27500			在整个种子发育过程中维持高表达		正向调控种子休眠	降解胚细胞间的胼胝质, 调控发育种子中ABA的积累	拟南芥		Wang et al., 2023
SPT	AT4G36930			-		存在正向和反向两种调控种子休眠机制	在Landsberg erecta与Columbia中种子休眠的遗传表现相反。直接抑制RGA和MFT的表达, 直接调控ABI5的表达	拟南芥		Vaistij et al., 2013
OsDOR1	LOC_Os03g-20770			在种子胚中特异表达		正向调控种子休眠	OsDOR1与OsGID1互作, 破坏OsGID1-OsSLR1复合体的形成, 影响GA信号转导	水稻		Kim et al., 2023
FIP1	AT5G58040			在干种子中表达		正向调控种子休眠	FIP1是一个加工pre-mRNA 3'端的蛋白, Abi5、DOG1和PYL12在突变体fip1中表达量下降	拟南芥		Li et al., 2023
OsNCED3	LOC_Os03g-44380			在种子胚发育过程中高表达		反向调控种子休眠	突变后, 种子胚的ABA含量降低, GA含量升高; 而超表达植株中, 种子胚的ABA/GA比例关系到种子的休眠性	水稻		Chen et al., 2023
SFL1	AT1G27461			在种子发育过程中表达量逐渐升高		反向调控种子休眠	OsSdr4的同源基因, 且功能相同	拟南芥		Zheng et al., 2022
AtMLP329	AT2G01530			在种子胚的胚根中表达		正向调控种子的初级休眠	DOF6直接调控MLP329的表达; 突变后, GA合成酶基因GA1表达量升高, ABA合成酶基因ZEP表达量下降	拟南芥		Chong et al., 2022
TaETR1	TraesCS4A-02G274300			在所有组织中都有表达, 在根中表达量最高		正向调控种子休眠	超表达植株的种子对乙烯不敏感	小麦		Wei et al., 2023
AtAAH	AT4G20070			在发育中的角果、根和干种子及萌发期的种子中高表达		反向调控种子休眠	外施硝酸钾可部分恢复ataah种子的高休眠性表型	拟南芥		Yazdanpanah et al., 2022
OsABA8ox1	LOC_Os02g-47470			在种子中表达		反向调控种子休眠	降低ABA含量	水稻		Fu et al., 2022
OsABA8ox2	LOC_Os08g-36860
OsABA8ox3	LOC_Os09g-28390
KCS12			Mtr.49305.1.- S1_at		在种皮中表达	正向调控种子休眠	控制种皮细胞中长链脂肪酸的合成	蒺藜苜蓿		Chai et al., 2021
FHY3			AT3G22170		随着角果的发育, 表达量逐渐升高, 在干种子中表达量达到峰值	反向调控种子休眠	白光促进FHY3蛋白积累; FHY3与phyB互作, 直接调控RVE2和RVE7的表达, 直接抑制SPT的表达	拟南芥		Liu et al., 2021
FT			AT1G65480			正向调控种子休眠	在种子中特异表达FT或者TFL1, GA含量降低	拟南芥		Chen et al., 2021
TFL1			AT5G03840			正向调控种子休眠	在种子中特异表达FT或者TFL1, GA含量降低	拟南芥		Chen et al., 2021
OsbZIP09			LOC_Os01g-59760		ABA处理15分钟可诱导OsbZIP09的表达	反向调控种子休眠	利用RNA-seq和DAP-seq分析了52个OsbZIP09直接调控的基因, 包括休眠相关基因OsLOX2和LEA家族	水稻		Zhu et al., 2021
TaAMY2			TraesCS7D-02G380400		开花后5天高表达, 之后表达量逐渐降低	反向调控种子休眠	在超表达TaAmy2植株中, α-淀粉酶的活性升高, 导致可溶性糖含量升高, 新鲜种子没有休眠性, 对ABA不敏感	小麦		Zhang et al., 2021
OsGLP2-1			LOC_Os02g-29000		在种子盾片中特异表达	正向调控种子休眠	受ABA诱导表达, GA抑制其表达; ABI5和GAMYB拮抗地调控GLP2-1的表达	水稻		Wang et al., 2020a
REF6			AT3G48430		在发育中的角果表达	反向调控种子休眠	在角果发育过程中, REF6结合CYP707A1和CYP707A3, 并负责这2个基因的H3K27me3修饰	拟南芥		Chen et al., 2020b
HSFA9			Medtr4g126-070		在种子中特异表达	反向调控种子休眠	调控ABA代谢和信号通路	蒺藜苜蓿		Zinsmeis- ter et al., 2020
SD6			LOC_Os06g-06900		在种子发育过程中表达量逐渐降低	反向调控种子休眠	与OsICE2互作, 直接促进ABAOX3的表达	水稻		Xu et al., 2022
OsICE2			LOC_Os01g-70310		在种子发育过程中表达量逐渐升高	正向调控种子休眠	与SD6互作, 直接抑制ABAOX3的表达	水稻		Xu et al., 2022
OsDOG1L-3			LOC_Os01g-20030		在开花后15天的种子中表达量达到最大值, 后逐渐下降	正向调控种子休眠	正反馈影响ABA合成基因的表达, 增加种子中ABA含量	水稻		Wang et al., 2020b
OsbZIP75			LOC_Os09g-34060		在种子中高表达	正向调控种子休眠	OsbZIP75直接调控OsDOG1L-3	水稻
OsbZIP78			LOC_Os10g-38820		在种子中高表达	正向调控种子休眠	同OsbZIP75	水稻
OsBT1			LOC_Os02g-10800		在种子中特异表达, 开花后21天达到峰值; 在茎、叶、叶鞘和穗中几乎不表达	正向调控种子休眠	-	水稻		Song et al., 2020
AtPER1			AT1G48130		在种子中特异表达	正向调控种子的初级休眠	清除种子中的活性氧, 抑制ABA降解和促进GA合成	拟南芥		Chen et al., 2020a
ETR1/RDO3			AT1G66340		-	正向调控种子休眠	ERF12与TPL相互作用, 直接抑制DOG1的表达, 而ETR1是ERF12的上游基因, 参与调控ERF12的表达	拟南芥		Li et al., 2019
DOGL4			At4g18650		在新鲜种子和干种子中都有高表达; 在开花后6天胚乳中、开花后8天胚中开始表达	反向调控种子休眠	在F₁代种子的胚乳细胞中, 来自父本的基因甲基化DOG4的启动子, 降低其表达; 纯合突变体种子的休眠能力增强	拟南芥		Zhu et al., 2018
ROS1			AT2G36490		在干种子中表达	反向调控种子休眠	促进DOG4的表达	拟南芥		Zhu et al., 2018
GHNAC83		GlaUn057212		在叶片、花和根中高表达; 在球茎中低表达		正向调控种子休眠	GHNAC83直接抑制GHPP2C1的表达, 影响ABA信号; 直接结合GHIPT的启动子, 负调控CK的合成	唐菖蒲	Wu et al., 2019
GHPP2C		GlaUn078852		在根、叶、球茎、雄蕊、雌蕊和花瓣中都有表达		反向调控种子休眠	识别ABA分子	唐菖蒲	Wu et al., 2019
ASPG1		AT3G18490		在圆锥花序及萌发的种子中表达		反向调控种子休眠	可能通过影响GA信号通路, 影响种子休眠及萌发	拟南芥	Shen et al., 2018
EBS		AT4G22140		在种子发育早期及吸胀前24小时的种子中表达		正向调控种子休眠	植物特有的一类转录调节蛋白, 与SHL在调控种子休眠上功能冗余, 并且与AGL67互作	拟南芥	Narro-Die- go et al., 2017
GATA12		AT5G25830		在新鲜种子中表达量最高, 在干种子中也有表达		正向调控种子休眠	RGL12与DOF6互作, 直接调控GATA12的表达	拟南芥	Ravindran et al., 2017
KNOX4		Medtr5g011-070		在种皮细胞中显著表达		反向调控种子休眠	突变体种子的种皮角质层发生变化, 直接调控角质层合成基因的表达	蒺藜苜蓿	Chai et al., 2016
OsGA20ox2		LOC_Os01g- 66100		在发育中的种子和胚细胞中表达		反向调控种子休眠	影响发育中种子的GA合成基因表达	水稻	Ye et al., 2015
RDO5		AT4G11040		在种子中特异表达, 在干种子中表达量最高		正向调控种子休眠	RDO5编码一个PP2C磷酸酶, 通过抑制RNA结合蛋白APUM9的表达调控种子休眠	拟南芥	Xiang et al., 2014
APUM9		AT1G35730		在开花后16天的种子中高表达, 之后表达量逐渐降低		反向调控种子休眠	RDO5编码一个PP2C磷酸酶, 通过抑制RNA结合蛋白APUM9的表达调控种子休眠	拟南芥	Xiang et al., 2014
WRKY41		AT4g11070		在成熟种子、种子胚、叶脉和下胚轴中表达		正向调控种子休眠	WRKY14直接调控ABI3的表达, 但与ABA通路无关联	拟南芥	Ding et al., 2014
HON		AT1g07430		在开花后9天的种子中表达, 开花后12天表达量达到峰值		反向调控种子休眠	是PP2C家族蛋白, 在ABA存在时, 与PYR1/RCAR11互作; 抑制ABA信号而激活GA信号	拟南芥	Kim et al., 2013
ABI4		AT2G40220		在种子中高表达		反向调控种子休眠	ABI4直接抑制CYP707A1和CYO- 707A2的表达; 突变体中ABA含量降低, GA含量升高, abi4突变体可以恢复ga1-t不发芽的表型	拟南芥	Shu et al., 2013
SNL1		AT3G01320		随着种子成熟, 表达量逐渐升高		正向调控种子休眠	SNL1与SNL2形成组蛋白去乙酰化复合体, SNL1与组蛋白去乙酰化酶HDA19互作, 调控乙烯合成基因及ABA合成、降解基因的乙酰化水平	拟南芥	Wang et al., 2013
SNL2		AT5G15020		随着种子成熟, 表达量逐渐升高		正向调控种子休眠		拟南芥	Wang et al., 2013
AtDOF6		AT3G45610		在新鲜种子中表达, 在后熟过程和吸胀时期逐渐降低		正向调控种子休眠	与TCP14互作, TCP14反向调控种子休眠	拟南芥	Rueda-Ro mero et al., 2012
AtET2		AT5G56780		在成熟的种子胚中表达		正向调控种子休眠	FUS3直接抑制AtET2的表达	拟南芥	Ivanov et al., 2012
FUS3		AT3G26790		在种子胚发育过程中持续表达, 在成熟种子中高表达		反向调控种子休眠	FUS3直接抑制AtET2的表达	拟南芥	Ivanov et al., 2012
KYP/SUVH4		AT5G13960		在吸胀种子中表达量最高		反向调控种子休眠	突变体中, DOG1和ABI3的表达量升高	拟南芥	Zheng et al., 2012
AMP1			AT3G54720	在种子发育过程中表达, 在开花后8天的种子中表达量最高		反向调控种子休眠	在不同背景的拟南芥中, 突变后的种子休眠表型不一致, 初步鉴定突变后影响ABA含量	拟南芥		Griffiths et al., 2011
TaMFT			AB456688	在不成熟胚的盾片和胚芽鞘中表达; 在种子发育过程中保持高表达		正向调控种子休眠	AtMFT的同源基因	小麦		Nakamu- ra et al., 2011
NCED6			AT3G24220	在发育中的种子胚乳中表达		正向调控种子休眠	在种子中诱导表达NCED6后, 种子中的ABA含量升高, 萌发率下降, 新鲜种子的休眠率升高	拟南芥		Martínez-Andújar et al., 2011
DOG1			AT5G45830	仅在种子中表达, 且在开花后9天表达量最高		正向调控种子休眠	-	拟南芥		Bentsink et al., 2006

基因名称	ID			表达特点		功能	分子机制	物种		参考文献
SDR3.1	LOC_Os03g-11550			开花后5天开始表达, 15天达到峰值		反向调控种子休眠	与ABI5互作, 并抑制ABI5的表达	水稻		Guo et al., 2024
SRO1	TraesCS4A-01G321300			在叶片、根和种子中表达		反向调控种子休眠	与TaVP1互作, 抑制穗上萌发基因TaPHS1和TaSdr的表达, 且影响TaVP1与TaABI5的互作	小麦		Liu et al., 2024
OsWSD1	LOC_Os03g-24460			在种子发育过程中, 其表达量逐渐降低; 在种子吸胀前36小时稳定表达		正向调控种子休眠	抑制GA的合成和α-淀粉酶的活性; 突变后对ABA不敏感	水稻		Huang et al., 2023
AtAIL6	AT5G10510			在整个种子发育过程中维持高表达		正向调控种子休眠	FUS3直接调控AtAIL6的表达	拟南芥		Liu et al., 2023
WRKY36	AT1G69810			在种子中高表达		反向调控种子休眠	WRKY36与AFP2互作, 复合体直接抑制DOG1的表达	拟南芥		Deng et al., 2023
AFP2	AT1G13740			在种子中高表达		反向调控种子休眠	WRKY36与AFP2互作, 复合体直接抑制DOG1的表达	拟南芥		Deng et al., 2023
OsNAC2	LOC_Os04g-38720			在种子发育和萌发过程中高表达		正向调控种子休眠	直接抑制OsABAox1和OsABAox2的表达	水稻		Zhao et al., 2023
BG14	AT2G27500			在整个种子发育过程中维持高表达		正向调控种子休眠	降解胚细胞间的胼胝质, 调控发育种子中ABA的积累	拟南芥		Wang et al., 2023
SPT	AT4G36930			-		存在正向和反向两种调控种子休眠机制	在Landsberg erecta与Columbia中种子休眠的遗传表现相反。直接抑制RGA和MFT的表达, 直接调控ABI5的表达	拟南芥		Vaistij et al., 2013
OsDOR1	LOC_Os03g-20770			在种子胚中特异表达		正向调控种子休眠	OsDOR1与OsGID1互作, 破坏OsGID1-OsSLR1复合体的形成, 影响GA信号转导	水稻		Kim et al., 2023
FIP1	AT5G58040			在干种子中表达		正向调控种子休眠	FIP1是一个加工pre-mRNA 3'端的蛋白, Abi5、DOG1和PYL12在突变体fip1中表达量下降	拟南芥		Li et al., 2023
OsNCED3	LOC_Os03g-44380			在种子胚发育过程中高表达		反向调控种子休眠	突变后, 种子胚的ABA含量降低, GA含量升高; 而超表达植株中, 种子胚的ABA/GA比例关系到种子的休眠性	水稻		Chen et al., 2023
SFL1	AT1G27461			在种子发育过程中表达量逐渐升高		反向调控种子休眠	OsSdr4的同源基因, 且功能相同	拟南芥		Zheng et al., 2022
AtMLP329	AT2G01530			在种子胚的胚根中表达		正向调控种子的初级休眠	DOF6直接调控MLP329的表达; 突变后, GA合成酶基因GA1表达量升高, ABA合成酶基因ZEP表达量下降	拟南芥		Chong et al., 2022
TaETR1	TraesCS4A-02G274300			在所有组织中都有表达, 在根中表达量最高		正向调控种子休眠	超表达植株的种子对乙烯不敏感	小麦		Wei et al., 2023
AtAAH	AT4G20070			在发育中的角果、根和干种子及萌发期的种子中高表达		反向调控种子休眠	外施硝酸钾可部分恢复ataah种子的高休眠性表型	拟南芥		Yazdanpanah et al., 2022
OsABA8ox1	LOC_Os02g-47470			在种子中表达		反向调控种子休眠	降低ABA含量	水稻		Fu et al., 2022
OsABA8ox2	LOC_Os08g-36860
OsABA8ox3	LOC_Os09g-28390
KCS12			Mtr.49305.1.- S1_at		在种皮中表达	正向调控种子休眠	控制种皮细胞中长链脂肪酸的合成	蒺藜苜蓿		Chai et al., 2021
FHY3			AT3G22170		随着角果的发育, 表达量逐渐升高, 在干种子中表达量达到峰值	反向调控种子休眠	白光促进FHY3蛋白积累; FHY3与phyB互作, 直接调控RVE2和RVE7的表达, 直接抑制SPT的表达	拟南芥		Liu et al., 2021
FT			AT1G65480			正向调控种子休眠	在种子中特异表达FT或者TFL1, GA含量降低	拟南芥		Chen et al., 2021
TFL1			AT5G03840			正向调控种子休眠	在种子中特异表达FT或者TFL1, GA含量降低	拟南芥		Chen et al., 2021
OsbZIP09			LOC_Os01g-59760		ABA处理15分钟可诱导OsbZIP09的表达	反向调控种子休眠	利用RNA-seq和DAP-seq分析了52个OsbZIP09直接调控的基因, 包括休眠相关基因OsLOX2和LEA家族	水稻		Zhu et al., 2021
TaAMY2			TraesCS7D-02G380400		开花后5天高表达, 之后表达量逐渐降低	反向调控种子休眠	在超表达TaAmy2植株中, α-淀粉酶的活性升高, 导致可溶性糖含量升高, 新鲜种子没有休眠性, 对ABA不敏感	小麦		Zhang et al., 2021
OsGLP2-1			LOC_Os02g-29000		在种子盾片中特异表达	正向调控种子休眠	受ABA诱导表达, GA抑制其表达; ABI5和GAMYB拮抗地调控GLP2-1的表达	水稻		Wang et al., 2020a
REF6			AT3G48430		在发育中的角果表达	反向调控种子休眠	在角果发育过程中, REF6结合CYP707A1和CYP707A3, 并负责这2个基因的H3K27me3修饰	拟南芥		Chen et al., 2020b
HSFA9			Medtr4g126-070		在种子中特异表达	反向调控种子休眠	调控ABA代谢和信号通路	蒺藜苜蓿		Zinsmeis- ter et al., 2020
SD6			LOC_Os06g-06900		在种子发育过程中表达量逐渐降低	反向调控种子休眠	与OsICE2互作, 直接促进ABAOX3的表达	水稻		Xu et al., 2022
OsICE2			LOC_Os01g-70310		在种子发育过程中表达量逐渐升高	正向调控种子休眠	与SD6互作, 直接抑制ABAOX3的表达	水稻		Xu et al., 2022
OsDOG1L-3			LOC_Os01g-20030		在开花后15天的种子中表达量达到最大值, 后逐渐下降	正向调控种子休眠	正反馈影响ABA合成基因的表达, 增加种子中ABA含量	水稻		Wang et al., 2020b
OsbZIP75			LOC_Os09g-34060		在种子中高表达	正向调控种子休眠	OsbZIP75直接调控OsDOG1L-3	水稻
OsbZIP78			LOC_Os10g-38820		在种子中高表达	正向调控种子休眠	同OsbZIP75	水稻
OsBT1			LOC_Os02g-10800		在种子中特异表达, 开花后21天达到峰值; 在茎、叶、叶鞘和穗中几乎不表达	正向调控种子休眠	-	水稻		Song et al., 2020
AtPER1			AT1G48130		在种子中特异表达	正向调控种子的初级休眠	清除种子中的活性氧, 抑制ABA降解和促进GA合成	拟南芥		Chen et al., 2020a
ETR1/RDO3			AT1G66340		-	正向调控种子休眠	ERF12与TPL相互作用, 直接抑制DOG1的表达, 而ETR1是ERF12的上游基因, 参与调控ERF12的表达	拟南芥		Li et al., 2019
DOGL4			At4g18650		在新鲜种子和干种子中都有高表达; 在开花后6天胚乳中、开花后8天胚中开始表达	反向调控种子休眠	在F₁代种子的胚乳细胞中, 来自父本的基因甲基化DOG4的启动子, 降低其表达; 纯合突变体种子的休眠能力增强	拟南芥		Zhu et al., 2018
ROS1			AT2G36490		在干种子中表达	反向调控种子休眠	促进DOG4的表达	拟南芥		Zhu et al., 2018
GHNAC83		GlaUn057212		在叶片、花和根中高表达; 在球茎中低表达		正向调控种子休眠	GHNAC83直接抑制GHPP2C1的表达, 影响ABA信号; 直接结合GHIPT的启动子, 负调控CK的合成	唐菖蒲	Wu et al., 2019
GHPP2C		GlaUn078852		在根、叶、球茎、雄蕊、雌蕊和花瓣中都有表达		反向调控种子休眠	识别ABA分子	唐菖蒲	Wu et al., 2019
ASPG1		AT3G18490		在圆锥花序及萌发的种子中表达		反向调控种子休眠	可能通过影响GA信号通路, 影响种子休眠及萌发	拟南芥	Shen et al., 2018
EBS		AT4G22140		在种子发育早期及吸胀前24小时的种子中表达		正向调控种子休眠	植物特有的一类转录调节蛋白, 与SHL在调控种子休眠上功能冗余, 并且与AGL67互作	拟南芥	Narro-Die- go et al., 2017
GATA12		AT5G25830		在新鲜种子中表达量最高, 在干种子中也有表达		正向调控种子休眠	RGL12与DOF6互作, 直接调控GATA12的表达	拟南芥	Ravindran et al., 2017
KNOX4		Medtr5g011-070		在种皮细胞中显著表达		反向调控种子休眠	突变体种子的种皮角质层发生变化, 直接调控角质层合成基因的表达	蒺藜苜蓿	Chai et al., 2016
OsGA20ox2		LOC_Os01g- 66100		在发育中的种子和胚细胞中表达		反向调控种子休眠	影响发育中种子的GA合成基因表达	水稻	Ye et al., 2015
RDO5		AT4G11040		在种子中特异表达, 在干种子中表达量最高		正向调控种子休眠	RDO5编码一个PP2C磷酸酶, 通过抑制RNA结合蛋白APUM9的表达调控种子休眠	拟南芥	Xiang et al., 2014
APUM9		AT1G35730		在开花后16天的种子中高表达, 之后表达量逐渐降低		反向调控种子休眠	RDO5编码一个PP2C磷酸酶, 通过抑制RNA结合蛋白APUM9的表达调控种子休眠	拟南芥	Xiang et al., 2014
WRKY41		AT4g11070		在成熟种子、种子胚、叶脉和下胚轴中表达		正向调控种子休眠	WRKY14直接调控ABI3的表达, 但与ABA通路无关联	拟南芥	Ding et al., 2014
HON		AT1g07430		在开花后9天的种子中表达, 开花后12天表达量达到峰值		反向调控种子休眠	是PP2C家族蛋白, 在ABA存在时, 与PYR1/RCAR11互作; 抑制ABA信号而激活GA信号	拟南芥	Kim et al., 2013
ABI4		AT2G40220		在种子中高表达		反向调控种子休眠	ABI4直接抑制CYP707A1和CYO- 707A2的表达; 突变体中ABA含量降低, GA含量升高, abi4突变体可以恢复ga1-t不发芽的表型	拟南芥	Shu et al., 2013
SNL1		AT3G01320		随着种子成熟, 表达量逐渐升高		正向调控种子休眠	SNL1与SNL2形成组蛋白去乙酰化复合体, SNL1与组蛋白去乙酰化酶HDA19互作, 调控乙烯合成基因及ABA合成、降解基因的乙酰化水平	拟南芥	Wang et al., 2013
SNL2		AT5G15020		随着种子成熟, 表达量逐渐升高		正向调控种子休眠		拟南芥	Wang et al., 2013
AtDOF6		AT3G45610		在新鲜种子中表达, 在后熟过程和吸胀时期逐渐降低		正向调控种子休眠	与TCP14互作, TCP14反向调控种子休眠	拟南芥	Rueda-Ro mero et al., 2012
AtET2		AT5G56780		在成熟的种子胚中表达		正向调控种子休眠	FUS3直接抑制AtET2的表达	拟南芥	Ivanov et al., 2012
FUS3		AT3G26790		在种子胚发育过程中持续表达, 在成熟种子中高表达		反向调控种子休眠	FUS3直接抑制AtET2的表达	拟南芥	Ivanov et al., 2012
KYP/SUVH4		AT5G13960		在吸胀种子中表达量最高		反向调控种子休眠	突变体中, DOG1和ABI3的表达量升高	拟南芥	Zheng et al., 2012
AMP1			AT3G54720	在种子发育过程中表达, 在开花后8天的种子中表达量最高		反向调控种子休眠	在不同背景的拟南芥中, 突变后的种子休眠表型不一致, 初步鉴定突变后影响ABA含量	拟南芥		Griffiths et al., 2011
TaMFT			AB456688	在不成熟胚的盾片和胚芽鞘中表达; 在种子发育过程中保持高表达		正向调控种子休眠	AtMFT的同源基因	小麦		Nakamu- ra et al., 2011
NCED6			AT3G24220	在发育中的种子胚乳中表达		正向调控种子休眠	在种子中诱导表达NCED6后, 种子中的ABA含量升高, 萌发率下降, 新鲜种子的休眠率升高	拟南芥		Martínez-Andújar et al., 2011
DOG1			AT5G45830	仅在种子中表达, 且在开花后9天表达量最高		正向调控种子休眠	-	拟南芥		Bentsink et al., 2006

重要的种子储存物质长寿命mRNA

Indispensable Material for Germination: Long-lived mRNAs of Plant Seed

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