林木种子休眠与萌发机制研究进展

doi:10.11707/j.1001-7488.20211013

摘要/Abstract

摘要：

种子休眠是由外界环境因素或者种子自身遗传特性诱导产生的，是具有生活力的种子在有利环境下仍不能萌发的现象。种子休眠与萌发是2种既相互区别又相互联系的发育状态，具有休眠特性的种子只有休眠解除才可能萌发，种子休眠在一定程度上决定其发芽所需的环境。为了对种子休眠与萌发现象有更全面的认识，本文在收集国内外林木种子休眠、休眠解除及萌发研究文献的基础上，总结了常见林木种子休眠产生的原因以及休眠类型划分系统，探讨休眠产生的机制，阐述内源激素调控种子休眠、阐述光信号调控种子休眠、Ca²⁺调控种子休眠、基因控制种子休眠、细胞膜变论等学说；概述了种子通过层积处理、物理方法、化学方法以及激素等解除休眠的方法；种子恢复吸胀则表示已进入萌发阶段，其伴随着形态、生理生化及分子层面的复杂变化。同时讨论当前研究的薄弱之处，并提出应对种子休眠与萌发的启动转录、基因调控、遗传表达等分子调控机制展开更深入的研究，以期为林木种子休眠解除与萌发机制方面的研究提供依据，为林木种子贮存、苗木繁育、种植推广及开发利用等提供参考。

关键词: 种子休眠, 影响因素, 调控机理, 休眠解除, 种子萌发

Abstract:

Seed dormancy is induced by external environmental factors or seed genetic characteristics, and it is a phenomenon that the viable seeds cannot germinate under a favorable environment. Seed dormancy and germination are two different and interrelated developmental states. Dormant seeds can enter into the germination state only after the dormancy isreleased. Seed dormancy to a certain extent determines the environment required for germination. In order to have a more comprehensive understanding ofseed dormancy and germination, this paper summarizes the causes of common forest seed dormancy and the classification system of the dormancy types based on the domestic and foreign literature regarding forest seed dormancy, dormancy release, and germination. Furthermore, in this paper we discuss the mechanism of seed dormancy, especially the theory of endogenous hormone regulating seed dormancy, expound the theories of light signal-regulating seed dormancy, Ca²⁺ regulating seed dormancy and gene controlling seed dormancy, cell membrane changes, etc; The methods of seed dormancy release by stratification, physical methods, chemical methods, and hormone treatments are summerized. The recovery of imbibition indicates that the seed has entered the germination stage, which is accompanied by complex changes in morphology, physiology, biochemistry, and molecular level. Moreover, this paper also discusses the limitation of current research and proposes that more in-depth research should be carried out on the molecular regulation mechanisms of seed dormancy and germination, such as initiation of transcription, gene regulation, genetic expression, in order to provide a basis for the study on dormancy and germination mechanism of forest seeds and provide a reference for forest seed storage, seedling breeding, planting promotion and development, and utilization.

Key words: seed dormancy, influencing factors, regulatory mechanism, dormancy release, seed germination

中图分类号:

王艳梅,张小雪,朱秀征,张志华,李志,耿晓东,蔡齐飞,刘震. 林木种子休眠与萌发机制研究进展[J]. 林业科学, 2021, 57(10): 128-144.

Yanmei Wang,Xiaoxue Zhang,Xiuzheng Zhu,Zhihua Zhang,Zhi Li,Xiaodong Geng,Qifei Cai,Zhen Liu. Research Progress on Dormancy and Germination Mechanism of Forest Seeds[J]. Scientia Silvae Sinicae, 2021, 57(10): 128-144.

图/表 7

表1

脱落酸代谢及信号通路中与种子休眠调控相关基因"

关键基因Key gene	功能Function	通路中位置Location in pathway	参考文献Reference
ABF1，ABF2	在信号通路中起负调控作用 A negative regulatory role in the signaling pathway	抑制因子 Inhibitory factor	Ma et al., 2009;
HONSU	过表达则导致浅种子休眠，其突变体表现出较深的休眠 Overexpression led to shallow seed dormancy and its mutants showed deep dormancy	抑制因子 Inhibitory factor	Kim et al., 2013
AHG1，AHG3	ABA信号传递和种子休眠的负调控因子，AHG1或AHG3的功能缺失将增强种子休眠 Negative regulators of ABA signaling and seed dormancy. The loss of AHG1 or AHG3 will enhance seed dormancy	抑制因子 Inhibitory factor	NÉE et al., 2017
DOG1	通过抑制脱落酸通路的负调控因子AHG1的磷酸酶活性促进休眠 Promoting dormancy by inhibiting the phosphatase activity of AHG1, a negative regulator of the abscisic acid pathway	抑制因子 Inhibitory factor	Noriyuki et al., 2018
EDP	影响ABA敏感性 Affecting the sensitivity of ABA	转录因子 Transcription factor	Barrero et al., 2010
MYB96	调控ABI4以及ABA合成相关基因的转录进而促进休眠 Regulation the related genes of transcription of ABI4 and ABA synthesis to promote dormancy	转录因子 Transcription factor	Lee et al., 2015a 刘晏等，2020；
MYB33，MYBl01	调控ABA的含量及其表达水平，从而引起种子的休眠与成熟 Regulation the content and expression level of ABA, thus causing seed dormancy and maturity	转录因子 Transcription factor	Reyes et al., 2007
WRKY6	直接结合关键靶基因RAV1的启动子抑制其表达, 进而调控ABA Binding to the promoter of the key target gene RAV1 to inhibit its expression and regulate ABA	转录因子 Transcription factor	Huang et al., 2016
ARF10，ARF16	调控ABI3的表达，从而维持ABA信号的强度，进而促进种子休眠 Regulation the expression of ABI3, the intensity of ABA signal can be maintained and seed dormancy can be promoted	转录因子 Transcription factor	帅海威等，2016
DOG1	能与AHG1/AHG3结合，通过结合ABA信号传递的负调控因子和增加对ABA的敏感性而引起种子休眠。 Combining with AHG1/AHG3 and induce seed dormancy by combining negative regulatory factors of ABA signaling and increasing sensitivity to ABA	转录因子 Transcription factor	宋松泉等，2020a
HUB1	HUB1的消除导致ABA代谢和反应相关基因；HUB1的突变会导致种子休眠的减少 The elimination of HUB1 will lead to ABA metabolism and response-related genes；the mutation of HUB1 will lead to the decrease of seed dormancy	转录因子 Transcription factor
EGE1	ABA超敏突变体，能够增强休眠 ABA hypersensitive mutants can enhance dormancy	突变体 Mutant	Liu et al., 2007
HDA19	功能缺失突变体Hda19-1和Hda19-2都显示种子休眠减少 Both Hda19-1 and Hda19-2 showed reduced seed dormancy	突变体 Mutant	Long et al., 2006；Chen et al., 2010
NCED6，NCED9	促进ABA的积累和诱导休眠的作用 Promoting the accumulation of ABA and induce dormancy	突变体 Mutant	Valérie et al., 2010
CYP707A3，NCED4	ABA生物合成的关键基因，在休眠维持和解除阶段的表达水平上调 The key gene of ABA biosynthesis is up-regulated in the stage of dormancy maintenance and release	突变体 Mutant	Jia et al., 2020
KYP/SUVH4	在种子中的过度表达导致休眠和ABA敏感性降低Overexpression in seeds leads to dormancy and decreased ABA sensitivity	突变体Mutant
SUVH4，SUVH5	抑制DOG1和ABI3转录Inhibition of DOG1 and ABI3 transcription	突变体Mutant	Jian et al., 2012
BIN2	磷酸化并稳定ABI5以增强ABA信号 Phosphorylation and stabilization of ABI5 to enhance ABA signal	突变体 Mutant	Hu et al., 2014
LDL1，LDL2	通过负调控DOG1抑制种子休眠 Inhibition of seed dormancy by negative regulation of DOG1	突变体 Mutant	Zhao et al., 2015
DEP	促进ABI3转录 Promoting ABI3 transcription	突变体 Mutant	Lee et al., 2015b
Oswrky29	下调OsABF1、OsVP1表达抑制休眠 Inhibition of dormancy by downregulating the expression of OsABF1 and OsVP1	突变体 Mutant	Zhou et al., 2020
SNL1，SNL2	ABA途径的正调节因子 Positive regulators of the ABA pathway	蛋白 Protein	Wang et al., 2013
VQ18，VQ26	作为ABI5转录因子的抑制子负调控ABA信号转导 As an inhibitor of the ABI5 transcription factor, it negatively regulates ABA signal transduction	蛋白 Protein	Pan et al., 2018
RGL2	DELLA蛋白，通过与转录因子NF-YC互作, 诱导下游ABI5的转录 DELLA protein by interacting with transcription factor NF-YC induces the transcription of downstream ABI5	蛋白 Protein	Liu et al., 2016
CYP707A	突变体导致休眠程度加深 Mutants lead to deeper dormancy	代谢酶 Metabolizing enzyme	Gonzalez et al., 2004
CPK4，CPK11	作为正调控因子参与ABA信号途径 As a positive regulator involved in ABA signaling pathway	激酶 Kinase	Zhu et al., 2007
CPK12	负调控ABA信号途径 Negative regulation of ABA signaling pathway	激酶 Kinase	Rui et al., 2011
RAF10，RAF11	影响ABI3和ABI5转录来调节种子休眠 Regulating seed dormancy by affecting ABI3 and ABI5 transcription	激酶 Kinase	Lee et al., 2015b

表1

表2

赤霉素代谢及信号通路中与种子休眠调控相关基因"

关键基因 Key gene	功能 Function	通路中位置 Location in access	参考文献 Reference
CTS	可能作为专一的GA信号组分提高种子发育的潜能，从而打破胚的休眠 May act as a specific GA signal component to enhance the potential of seed development and break embryo dormancy	突变体 Mutant	Russell et al., 2000
GA1，GA2	GA缺陷型突变体，显示出强烈的种子休眠 GA deficient mutant showed strong seed dormancy	突变体 Mutant	Lee et al., 2002
DDF1	直接促进GA2ox7，从而降低GA含量 Directly promote the growth of GA2ox7 and reduce the content of GA	突变体 Mutant	Magome et al., 2008
PIF1	抑制GA合成相关基因的表达，促进GA代谢相关基因的表达，进而下调内源GA₄的水平 Inhibiting the expression of GA synthesis related genes, promote the expression of GA metabolism-related genes, and then down-regulate the level of endogenous GA₄	蛋白 Protein	Oh et al., 2006
GAI	DELLA蛋白，能够通过抑制TCP14和TCP15的表达减缓细胞周期的进程，最终使种子保持休眠状态 DELLA protein can slow down the process of the cell cycle by inhibiting the expression of TCP14 and TCP15, and finally make the seeds remain dormant	蛋白 Protein	Resentini et al., 2015
DOG1	调控响应GA信号的细胞壁重塑相关基因的表达促进种子休眠 Regulating the expression of cell wall remodeling related genes in response to GA signaling to promote seed dormancy	转录因子 Transcription factor	Graeberet al., 2014；宋松泉等，2020b
CYP72A9	参与初生休眠，调节种子生物活性GA₄的稳态 Participating in primary dormancy and regulates the homeostasis of GA₄	转录因子 Transcription factor	He et al., 2019
AtSdr4L	可能通过介导DOG1、GA途径调节种子的休眠 May regulate seed dormancy by DOG1 and GA pathway	转录因子 Transcription factor	刘晏等，2020

表2

表3

生长素代谢及信号通路中与种子休眠调控相关基因"

关键基因 Key gene	功能 Function	通路中所在位置 Location in access	参考文献 Reference
YUC1，YUC6	yuc1 yuc6双突变体被用来研究种子休眠特性的变化 yuc1 and yuc6 double mutants were used to study the changes of seed dormancy characteristics	合成酶 Synthetase	Liu et al., 2013
IAA7/AXR2，IAA17/AXR3	其功能获得型突变体种子的休眠水平降低 The dormancy level of the functional mutant seeds decreased	抑制因子 Inhibitory factor	Chapman，2009；Liu et al., 2013
ARP，DRM	与IAA代谢相关；ARP和DMR1受ABA代谢调控，与休眠进程密切相关 Regulated by IAA metabolism; ARP and DMR1 are regulated by ABA metabolism and are closely related to the dormancy process	抑制因子 Inhibitory factor	Footitt et al., 2011
AXR2，AXR3	生长素信号通路中的负调控因子 Negative regulators in the auxin signaling pathway	抑制因子 Inhibitory factor	Liu et al., 2013
AUX/IAA，AUX1，SAUR	生长素转录的负调控因子 Negative regulators of auxin transcription	受体 Recipient	帅海威等，2016
TIR1/AFBs	生长素受体；其三、四突变体表型非常强烈、不能正常发育；双突变体种子的休眠程度显著降低 Auxin receptor; the phenotype of the third and fourth mutants was very strong and could not develop normally; the dormancy degree of the double mutant seeds decreased significantly	受体 Recipient
ARF16	激活ABI3的表达，从而使种子保持休眠状态 Activating ABI3 expression and keep seeds dormant	转录因子 Transcription factor	Chapman et al., 2009；Liu et al., 2013
ARF2	生长素响应因子，与其靶标基因HB33可以调节ABA信号的输出 Auxin response factor and its target gene HB33 can regulate the output of ABA signal	转录因子 Transcription factor	Wang et al., 2011a
ARF10	过量表达能够明显增强ABI3、ABI4、ABI5以及NCED9的表达, 进而提高种子对ABA的敏感性，增强休眠性 Overexpression could significantly enhance the expression of ABI3, ABI4, ABI5, and NCED9, and further, improve the sensitivity of seeds to ABA and enhance dormancy	转录因子 Transcription factor	李继洋等，2018

表3

表4

表5

激素代谢及信号途径中关键基因与种子萌发调控"

植物激素 Plant hormone	生物学功能 Biological function	参考文献 Reference
Auxin	IAA30是非典型生长素转录抑制子，与典型转录抑制子有着相似作用；高盐环境诱导其转录，并进而抑制种子萌发 IAA30 is an atypical auxin transcriptional repressor, which has a similar effect to the typical one; A high salt environment induces its transcription and further inhibits seed germination	Remington et al.，2004；Park et al.，2011
	种子含TIR1，AFB2，AFB3和TIR1，AFB1，AFB2，AFB3的突变受体时，会阻碍种子对生长素信号的接收, 使得胚胎发育不良, 丧失萌发能力 When seeds contain the mutant receptors of TIR1, AFB2, AFB3 and TIR1, AFB1, AFB2, AFB3, it will block the reception of auxin signal, resulting in poor embryo development and loss of germination ability	Liu et al.，2013；帅海威等，2016
	AUX1是ABA抑制种子萌发所必需的，其功能缺失突变体表现出对ABA的抗性增加 AUX1 is necessary for ABA to inhibit seed germination, and its function deficient mutants show increased resistance to ABA	Thole et al.，2014
	IAA反应因子(ARF2、ARF6和ARF19)和SAUR32，它们在胚胎发育过程中高度表达。 Auxin response factors (ARF2, ARF6, and ARF19) and SAUR32 are highly expressed during embryonic development	Jia et al.，2020
GA	DELLA蛋白GAI、RGA、RGL1、RGL2和RGL3，以及GA信号的负调节因子SPY通过维持休眠状态来抑制种子萌发活动 DELLA proteins GAI, RGA, RGL1, RGL2, and RGL3, and the negative regulator of GA signal, SPY, inhibit seed germination by maintaining dormancy	Lees et al.，2002；Davière et al., 2016
GA	DHGA12能与GA受体结合，也能促进种子萌发、下胚轴伸长和子叶变绿 DHGA12 can bind to GA receptors and promote seed germination, hypocotyl elongation, and cotyledon greening	Liu et al.，2019
ABA	转录因子HDA6/HDA19通过影响ABA诱导的基因表达来调节种子萌发 Transcription factors HDA6/HDA19 regulate seed germination by affecting ABA-induced gene expression	Long et al.，2006；Chen et al.，2010
ABA	通过在ABI5过表达中加入SnRK2蛋白激酶，可以重塑内源性ABI5磷酸化和抑制萌发 Adding SnRK2 protein kinase to the ABI5 overexpression line, endogenous ABI5 phosphorylation can be remolded and germination can be inhibited	Piskurewicz et al.，2008
SA	通过抑制GA诱导的α淀粉酶编码基因的表达抑制种子萌发 Inhibiting the expression of GA induced-α-amylase gene to inhibit seed germination	Xie et al.，2007
SA	在高盐胁迫下，SA通过其他通路促进种子萌发 Under high salt stress, SA promoted seed germination through other pathways	Lee et al.，2010
CTK	通过抑制ABI5的转录或促进ABI5蛋白的降解拮抗ABA效应 Antagonizing the ABA effect by inhibiting ABI5 transcription or promoting ABI5 protein degradation	Wang et al.，2011b；Guan et al.，2014
JA	JA通过抑制ABA合成相关基因的表达、促进ABA代谢相关基因的表达拮抗ABA，从而促进种子萌发 JA antagonizes ABA by inhibiting the expression of genes related to ABA synthesis and promoting the expression of genes related to ABA metabolism, thus promoting seed germination	Jacobsen et al.，2013
JA	转录因子JAZ3通过与ABI5互作抑制ABI5的转录激活活性，进而促进种子萌发 JAZ3, a transcription factor, inhibits ABI5 transcriptional activation by interacting with ABI5, thus promoting seed germination	Ju et al.，2019
BR	BR信号通路中的负调控因子BIN2，能够磷酸化并稳定ABI5蛋白，进而促进ABA信号转导 BIN2, a negative regulator of the BR signaling pathway, can phosphorylate and stabilize ABI5 protein, thereby promoting ABA signal transduction	Hu et al.，2014
BR	BRL2、BRL3、BEH3和BEH4，在早期胚胎发育阶段高度表达 BRL2, BRL3, BEH3, and BEH4 were highly expressed during early embryonic development	Jia et al.，2020

表5

表6

miRNA参与种子萌发调控功能"

miRNA	靶基因Gene of interest	生物学功能Biological function	参考文献Reference
miR167	ARF6，ARF8	生长素等植物激素信号传导 Signal transduction of auxin and other plant hormones	Xie et al.，2005
miR397	LAC	细胞壁加厚与木质化 Cell wall thickening and lignification	Luo et al.，2006
miR398		对细胞维持分裂具有重要作用 Playing an important role in maintaining cell division	Luo et al.，2006
miR402	DML3	调控种胚及胚乳发育 Regulation of embryo and endosperm development	Nonogaki，2010
miR1512		种皮形态建成 Seed coat morphogenesis	Zabala et al.，2012
miR166	HD-ZIPIII	细胞分裂分化 Cell division and differentiation	Zhang et al.，2013
miR167	ARF6，ARF8	生长素信号传导；种子形态建成 Auxin signal transduction and seed morphogenesis	Zabala et al.，2012；Gu et al.，2013；
miR5801	DME	DNA去甲基化；种胚及胚乳发育 DNA demethylation and embryo and endosperm development	Chio et al.，2002；Huang et al.，2013
miR5807	PPR	调控种皮、胚、胚乳发育 Regulating the development of seed coat, embryo, and endosperm	Huang et al.，2013；Liu et al.，2013
miR169	CCAAT-box	调控种胚形态建成 Regulation of embryo morphogenesis	Xing et al.，2017
miR160	ARF10，ARF10，ARF17	下调转录因子ARF10、ARF16和ARF17的表达，其异位表达导致其对ABA的敏感性降低 Decreasing the ectopic expression of ARF10, ARF16, and ARF17, and decreased their sensitivity to ABA	Wang et al.，2011；Curaba et al.，2014；Daset al.，2015；魏俊等，2017
miR390	ARF2	调控种子对ABA的敏感性 Regulation of seed sensitivity to ABA
miR159	MYB33，MYB101，MYB65	控制MYB转录因子蛋白转录水平，影响ABA敏感性；参与调控GA信号 Control the transcription level of MYB transcription factor protein and affect the sensitivity of ABA；Participate in regulating GA signal	Chenet al.，2012；Daset al.，2015
miR164	NAC	调控胚乳发育 Regulation of endosperm development	Li et al.，2016
miR163	PXMT1	光响应下促进种子萌发 Promoting seed germination under light response	Chung et al.，2016
miR393	TIR1，AFB2，AFB3	影响种子萌发对ABA的敏感性；调控胚乳发育 Affecting the sensitivity of seed germination to ABA; regulation of endosperm development	Zhu et al.，2017；魏俊等，2017；
miR171，miR2118		在发育的胚乳中高表达；参与胚乳中维生素生物合成与代谢过程 Highly expressed in the developing endosperm; It is involved in the biosynthesis and metabolism of vitamins in the endosperm	Zhu et al.，2017
miR156miR172	SPL13，SPL2AP2	两者交互作用，调控相关基因(如DOG1)，促进或延迟种子萌发 Interacting to regulate related genes (such as DOG1) to promote or delay seed germination	Nonogaki，2010；Das et al.，2015；Jia et al.，2020

表6

图1

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关键基因 Key gene	功能 Function	通路中所在位置 Location in access	参考文献 Reference
ETRL-1，EIN2	乙烯不敏感型突变体，加深种子休眠 ETH insensitive mutants deepen seed dormancy	突变体 Mutant	Nonogaki，2014
CTRL	乙烯超敏感型突变体，减弱种子休眠 ETH hypersensitive mutants weaken seed dormancy	突变体 Mutant	Nonogaki，2014
RDO3	休眠突变体，由乙烯受体ETR1突变功能缺失引起 Dormant mutant, caused by the loss of ethylene receptor ETR1 mutation function	突变体 Mutant	Li et al., 2019
erf12	负调控种子休眠 Negative regulation of seed dormancy	响应因子 Response factor	Li et al., 2019
ERF9，ERF105，ERF112	乙烯生物合成信号 ETH biosynthesis signal	响应因子 Response factor	Wang et al., 2013
SNL1，SNL2	通过负调控乙烯生物合成和反应促进种子休眠 Promoting seed dormancy through negative regulation of ethylene biosynthesis and response	蛋白 Protein	Wang et al., 2013
ACO1，ACO4，ACO5	乙烯代谢基因，并参与对抗ABA对胚乳盖弱化和胚乳破裂的抑制作用，这对萌发和休眠很重要 ETH metabolizing genes are involved in the inhibition of ABA on endosperm cap weakening and endosperm rupture, which are important for germination and dormancy	蛋白 Protein	Linkies et al., 2009 Wang et al., 2013
ETR1	促进DOG1的转录, 进而促进种子休眠Promoting the transcription of DOG1 and seed dormancy	蛋白Protein	Li et al., 2019

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