林木种子休眠与萌发机制研究进展

doi:10.11707/j.1001-7488.20211013

Abstract

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

CLC Number:

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.

Figures/Tables 7

Table 1

Metabolism of abscisic acid and signaling genes involved in seed dormancy regulation"

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

Table 1

Table 2

Gibberellin metabolism and signaling genes involved in seed dormancy regulation"

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

Table 2

Table 3

Auxin metabolism and signaling genes involved in seed dormancy regulation"

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

Table 3

Table 4

Table 5

Key genes in hormone metabolism, signaling pathway, and regulation of seed germination"

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

Table 5

Table 6

The role of miRNAs involved in seed germination"

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

Table 6

Fig.1

References 0

	安毅鹏, 侯潇棐, 黄一凡, 等. 西藏砂生槐种子休眠及解除方法研究. 高原农业, 2020, 4 (3): 245- 248.
	An Y P , Hou X F , Huang Y F , et al. Study on seed dormancy and release method of Sophora moorcroftiana in Tibet. Journal of Plateau Agriculture, 2020, 4 (3): 245- 248.
	曹运梅, 张建, 向云, 等. 铜钱树种子休眠特性及破眠技术研究. 种子, 2019, 38 (8): 73- 76. doi: 10.3969/j.issn.1005-2690.2019.08.063
	Cao Y M , Zhang J , Xiang Y , et al. Study on the dormancy characteristics and dormancy breaking techniques of Paliurus hemsleyanus Rehder seed. Seed, 2019, 38 (8): 73- 76. doi: 10.3969/j.issn.1005-2690.2019.08.063
	陈伟, 陈绍安, 李江, 等. 中泰南五味子种子休眠机理初探. 西部林业科学, 2020, 49 (2): 44- 56.
	Chen W , Chen S A , Li J , et al. A preliminary study on dormancy mechanism of Kadsura ananosma seeds. Journal of West China Forestry Science, 2020, 49 (2): 44- 56.
	陈伟, 马绍宾, 陈宏伟. 林木种子休眠及其研究方法概述. 林业调查规划, 2010, 35 (1): 31- 36. doi: 10.3969/j.issn.1671-3168.2010.01.008
	Chen W , Ma S B , Chen H W . Summarization on dormancy mechanism and research methods of forest tree seed. Forest Inventory and Planning, 2010, 35 (1): 31- 36. doi: 10.3969/j.issn.1671-3168.2010.01.008
	陈香波, 刘杨, 赵明水, 等. 极度濒危树种羊角槭的种胚发育与休眠解除. 林业科学, 2017, 53 (4): 65- 73.
	Chen X B , Liu Y , Zhao M S , et al. Embryo development and dormancy releasing of Acer yangjuechi, the extremely endangered plant. Scientia Silvae Sinicae, 2017, 53 (4): 68- 76.
	崔慧. 2017. 银杏种子休眠不同阶段的转录调控研究. 扬州: 扬州大学硕士学位论文.
	Cui H. 2017. Transcriptional regulation of Ginkgo biloba seeds at different dormancy stages. Yangzhou: MS thesis of Yangzhou University. [in Chinese]
	段加玉, 勾天兵, 谢朋, 等. 温度对紫椴种子萌发及萌发过程中物质转化的影响. 中国农学通报, 2013, 29 (19): 31- 34. doi: 10.11924/j.issn.1000-6850.2013-0432
	Duan J Y , Gou T B , Xie P , et al. Effect of temperature on seed germination and temperature on substance transformation of stratified seeds of Tilia amurensis Rupr. Chinese Agricultural Science Bulletin, 2013, 29 (19): 31- 34. doi: 10.11924/j.issn.1000-6850.2013-0432
	范文丽. 2015. 长白楤木种子打破休眠的生理生态因素研究. 沈阳: 沈阳农业大学硕士学位论文.
	Fan W L. 2005. Study on physiological and ecological factors of seed dormancy of Aralia continentalis Kitagawa. Shenyang: MS thesis of Shenyang Agricultural University. [in Chinese]
	房丽莎, 徐自恒, 刘震, 等. 山桐子果实发育过程中内含物、内源激素及光合特性的变化. 林业科学, 2020, 56 (11): 41- 52. doi: 10.11707/j.1001-7488.20201105
	Fang L S , Xu Z H , Liu Z , et al. Changes of contents, endogenous hormones and photosynthetic characteristics of Idesia polycarpa fruit at the different developmental stages. Scientia Silvae Sinicae, 2020, 56 (11): 41- 52. doi: 10.11707/j.1001-7488.20201105
	方志荣, 苏智先, 胡进耀. 脱落酸、赤霉素和乙烯对种子休眠的萌发和调控. 西华师范大学学报: 自然科学版, 2007, 28 (2): 128- 132.
	Fang Z R , Su Z X , Hu J Y . Effects of abscisic acid, gibberellin and ethylene on seed dormancy. Journal of China West Normal University: Natural Sciences Edition, 2007, 28 (2): 128- 132.
	符近, 尤瑞麟, 顾增辉. 马占相思种子休眠的研究. 北京大学学报: 自然科学版, 1997, 33 (6): 79- 85.
	Fu J , You R L , Gu Z H . Study on seed dormancy of Acacia mangium. Acta Scientiarum Naturalium Universitatis Pekinensis, 1997, 33 (6): 79- 85.
	洑香香, 周晓东, 刘红娜. 2013. 山茱萸种子休眠机理与解除方法初探. 中南林业科技大学学报, 33(4): 7-12.
	Fu X X, Zhou X D, Liu H N. 2013. A preliminary study on seed dormancy mechanism and relieving methods in Cornus officinalis. Journal of Central South University of Forestry & Technology, 33(4): 7-12. [in Chinese]
	高博, 辛宇. 椴树种子休眠机理及催芽研究进展. 特种经济动植物, 2015, 18 (11): 37- 38. doi: 10.3969/j.issn.1001-4713.2015.11.020
	Gao B , Xin Y . Research progress on dormancy mechanism and germination of Tilia amurensis seeds. Special Economic Animal and Plant, 2015, 18 (11): 37- 38. doi: 10.3969/j.issn.1001-4713.2015.11.020
	贺桂兰. GA₃、H₂O₂对大叶女贞种子发芽性能的影响. 现代园艺, 2020, 43 (10): 5- 6. doi: 10.3969/j.issn.1006-4958.2020.10.002
	He G L . Effects of GA₃ and H₂O₂ on seed germination of Ligustrum lucidum. Modern Horticulture, 2020, 43 (10): 5- 6. doi: 10.3969/j.issn.1006-4958.2020.10.002
	何青松. 2018. 银杏种子休眠过程与休眠解除研究. 扬州: 扬州大学硕士学位论文.
	He Q S. 2018. Seed dormancy process and dormancy release in Ginkgo biloba. Yangzhou: MS thesis of Yangzhou University. [in Chinese]
	黄玉国, 王文章. 刺楸种子层积过程中内源植物激素的动态变化及激素在种子休眠中的作用. 植物研究, 1990, 10 (4): 121- 129.
	Huang Y G , Wang W Z . Dynamic changes in endogenous hormones in the seeds of Kalopanax septemlobus during stratification and roles of hormones in the seed dormancy. Bulletin of Botanical Research, 1990, 10 (4): 121- 129.
	姜宗庆, 李成忠, 周霞, 等. 紫楠种子休眠特性及解除措施研究. 种子, 2020, 39 (12): 92- 98. doi: 10.3969/j.issn.1005-2690.2020.12.064
	Jiang Z Q , Li C Z , Zhou X , et al. Study on dormancy charateristics and removal measures of Phoebe sheareri seeds. Seed, 2020, 39 (12): 92- 98. doi: 10.3969/j.issn.1005-2690.2020.12.064
	康云清. 植物激素对种子休眠和萌发调控机理的分析研究. 种子科技, 2016, 34 (9): 65. doi: 10.3969/j.issn.1005-2690.2016.09.043
	Kang Y Q . Study on the regulation mechanism of plant hormones on seed dormancy and germination. Seed Science and Technology, 2016, 34 (9): 65. doi: 10.3969/j.issn.1005-2690.2016.09.043
	柯碧英, 周春燕, 李金英, 等. 珍贵物种顶果木种子休眠及解除方法研究. 种子, 2020, 39 (2): 143- 147.
	Ke B Y , Zhou C Y , Li J Y , et al. Study on methods of seed dormancy and dormancy-release for rare species Acrocarpus fraxinfolius. Seed, 2020, 39 (2): 143- 147.
	郎文香, 李宏博, 孙丹, 等. 离体条件下TDZ对解除北五味子种子休眠的影响. 西北植物学报, 2010, 30 (9): 1871- 1875.
	Lang W X , Li H B , Sun D , et al. In vitro studies of TDZ on breakage of seed dormancy of Schisandra chinensis(Turcz) Baill. Acta Botanica Boreali-Occidentalia Sinica, 2010, 30 (9): 1871- 1875.
	李继洋, 代培红, 杨洁, 等. 基于ARF10转录因子功能验证及其对拟南芥种子休眠和萌发的影响. 新疆农业大学学报, 2018, 41 (2): 34- 41.
	Li J Y , Dai P H , Yang J , et al. Functional verification of transcription factor based on ARF10 and its effect on dormancy and germin ation of Arabidopsis thaliana seeds. Journal of Xinjiang Agricultural University, 2018, 41 (2): 34- 41.
	李蓉, 叶勇. 种子休眠与破眠机理研究进展. 西北植物学报, 2005, 25 (11): 2350- 2355. doi: 10.3321/j.issn:1000-4025.2005.11.038
	Li R , Ye Y . Progresses in seed dormancy and dormancy-breaking mechanisms. Acta Botanica Boreali-Occidentalia Sinica, 2005, 25 (11): 2350- 2355. doi: 10.3321/j.issn:1000-4025.2005.11.038
	李生平. 2004. 银杏种子萌发过程中贮藏物质代谢机理的研究. 南京: 南京林业大学硕士论文.
	Li S P. 2004. Studies on mechanism of storage substance metabolize of Gingko biloba seeds during germination. Nanjing: MS thesis of Nanjing Forestry University. [in Chinese]
	李宇赤, 王婉丽, 杨灵. CO₂激光处理对灯盏花种子萌发的影响研究. 种子, 2019, 38 (5): 114- 116. doi: 10.3969/j.issn.1005-2690.2019.05.096
	Li Y C , Wang W L , Yang L . Effect of CO₂ laser treatment on seed germination of Erigeron breviscapus. Seed, 2019, 38 (5): 114- 116. doi: 10.3969/j.issn.1005-2690.2019.05.096
	李振华, 徐如宏, 任明见, 等. 光敏色素感知光温信号调控种子休眠与萌发研究进展. 植物生理学报, 2019, 55 (5): 539- 546.
	Li Z H , Xu R H , Ren M J , et al. Advances in phytochrome regulating seed dormancy and germination by sensing light and temperature signals. Plant Physiology Journal, 2019, 55 (5): 539- 546.
	刘婷, 李娟, 张晓曼, 等. 不同浸种处理对美国无刺皂荚种子萌发的影响. 农业科学研究, 2018, 39 (4): 2- 4.
	Liu T , Li J , Zhang X M , et al. Study on germination characteristics of Gleditsia triacanthos var.inermis. Journal of Agricultural Sciences, 2018, 39 (4): 2- 4.
	刘序. 2019. 珊瑚朴种子休眠机理研究. 南京: 南京林业大学硕士学位论文, .
	Liu X. 2019. Studies on seed dormancy mechanism of Celtis julianae Schneid. Nanjing: MS thesis of Nanjing Forestry University. [in Chinese]
	刘晏, 李俊德, 李家儒. 脱落酸和赤霉素调控种子休眠与萌发研究进展. 生物资源, 2020, 42 (2): 157- 163.
	Liu Y , Li J D , Li J R . Review of investigation progress of ABA and GA regulating seed dormancy and germination. Biotic Resources, 2020, 42 (2): 157- 163.
	刘月洋. 2010. 天女木兰种子层积过程中物质变化及其与种子萌发的关系. 沈阳: 沈阳农业大学硕士学位论文.
	Liu Y Y. 2010. Study on the substance changes and their relationship with dormancy broken of M. sieboldii K. koch seeds during stratification. Shenyang: MS thesis of Shenyang Agricultural University. [in Chinese]
	刘震, 王艳梅, 蒋建平. 不同种源白花泡桐种子的休眠生理生态研究. 生态学报, 2004, 24 (5): 959- 964. doi: 10.3321/j.issn:1000-0933.2004.05.015
	Liu Z , Wang Y M , Jiang J P . Eco-physiological study on seed dormancy in Paulownia fortunei from different provenances. Acta Ecologica Sinica, 2004, 24 (5): 959- 964. doi: 10.3321/j.issn:1000-0933.2004.05.015
	龙汉利, 罗建勋, 辜云杰, 等. 桢楠种子萌发过程中抗氧化酶及贮藏物质的变化. 西南农业学报, 2013, 26 (3): 978- 981. doi: 10.3969/j.issn.1001-4829.2013.03.028
	Long H L , Luo J X , Gu Y J , et al. Changes of antioxidant enzymes and storage substance during Phoebe seed germination. Southwest China Journal of Agricultural Sciences, 2013, 26 (3): 978- 981. doi: 10.3969/j.issn.1001-4829.2013.03.028
	鲁强. 2019. 三桠苦种子休眠的解除方法和初步机理研究. 广州: 广州中医药大学博士学位论文.
	Lu Q. 2019. Study on the method and mechanism of releasing seed dormancy of Melicope pteleifolia. Guangzhou: PhD thesis of Guangzhou University of Traditional Chinese Medicine. [in Chinese]
	马拉青呼. 2019. 4个品种桧柏种子休眠特性及萌发生理研究. 呼和浩特: 内蒙古农业大学硕士学位论文.
	Ma L Q H. 2019. Study on dormancy characteristics and germination physiology of four varieties Sabina chinensis. seeds Cultivars. Hohhot: MS thesis of Inner Mongolia Agricultural University. [in Chinese]
	马天晓, 王艳梅, 李世荣, 等. 硫酸和低温处理对酸角种子发芽率的影响. 河南农业大学学报, 2012, 46 (2): 152- 155. doi: 10.3969/j.issn.1000-2340.2012.02.009
	Ma T X , Wang Y M , Li S R , et al. Effect of different treatments of germination rate of Tamarind seeds. Journal of Henan Agricultural University, 2012, 46 (2): 152- 155. doi: 10.3969/j.issn.1000-2340.2012.02.009
	潘琳, 徐程扬. 种子休眠与萌发过程的生理调控机理. 种子, 2016, 26 (6): 42- 47. doi: 10.3969/j.issn.1005-2690.2016.06.028
	Pan L , Xu C Y . Review on mechanisms of physiological modulation in the process of seed dormancy and germination. Seed, 2016, 26 (6): 42- 47. doi: 10.3969/j.issn.1005-2690.2016.06.028
	秦爱丽, 郭泉水, 马凡强, 等. 温度、光照和水分对珍稀濒危树种崖柏种子萌发的影响. 种子, 2020, 39 (2): 19- 24. doi: 10.3969/j.issn.1005-2690.2020.02.012
	Qin A L , Guo Q S , Ma F Q , et al. Effects of temperature, light and water conditions on seed germination of Thuja sutchuenensis Franch. Seed, 2020, 39 (2): 19- 24. doi: 10.3969/j.issn.1005-2690.2020.02.012
	帅海威, 孟永杰, 罗晓峰, 等. 生长素调控种子的休眠与萌发. 遗传, 2016, 38 (4): 314- 322.
	Shuai H W , Meng Y J , Luo X F , et al. The roles of auxin in seed dormancy and germination. Hereditas, 2016, 38 (4): 314- 322.
	宋松泉, 刘军, 徐恒恒, 等. 脱落酸代谢与信号传递及其调控种子休眠与萌发的分子机制. 中国农业科学, 2020a, 53 (5): 857- 873.
	Song S Q , Liu J , Xu H H , et al. ABA metabolism and signaling and their molecular mechanism regulating seed dormancy and germination. Scientia Agricultura Sinica, 2020a, 53 (5): 857- 873.
	宋松泉, 刘军, 黄荟, 等. 赤霉素代谢与信号转导及其调控种子萌发与休眠的分子机制. 中国科学: 生命科学, 2020b, 50 (6): 599- 615.
	Song S Q , Liu J , Huang H , et al. Gibberellin metabolism and signaling and its molecular mechanism in regulating seed germination and dormancy. Scientia Sinica Vitae, 2020b, 50 (6): 599- 615.
	宋松泉, 刘军, 杨华, 等. 细胞分裂素代谢与信号转导及其调控种子萌发与休眠的分子机制. 植物学报, 2021, 56 (2): 218- 231.
	Song S Q , Liu J , Yang H , et al. Cytokinin metabolism and signal transduction and the molecular mechanism of regulating seed germination and dormancy. Chinese Bulletin of Botany, 2021, 56 (2): 218- 231.
	藤伊正. 1980. 植物的休眠与发芽. 刘瑞征. 译. 北京: 科学出版社.
	Teng Y Z. 1980. Plant dormancy and germination. Liu R Z. Translated. Beijing: Science Press. [in Chinese]
	王贵禧, 梁丽松. 贮藏板栗休眠与萌芽的温度调控. 林业科学, 1999, 35 (3): 29- 33. doi: 10.3321/j.issn:1001-7488.1999.03.006
	Wang G X , Liang L S . Temperature regulation of dormancy and germination of Chestnut during storage. Scientia Silvae Sinicae, 1999, 35 (3): 29- 33. doi: 10.3321/j.issn:1001-7488.1999.03.006
	王艳华, 高述民, 李凤兰, 等. 大山樱种子休眠机理的探讨. 种子, 2005, 24 (5): 12- 16.
	Wang Y H , Gao S M , Li F L , et al. Discussion of dormancy mechanism of Prunus sargentill seeds. Seed, 2005, 24 (5): 12- 16.
	王艳梅, 刘震, 牛晓锋. 1年生泡桐不同部位顶芽内源激素的动态变化. 林业科学, 2012, 48 (7): 61- 65.
	Wang Y M , Liu Z , Niu X F . Dynamic changes of endogenous hormones in terminal buds from different crown position of one-year-old Paulownia. Scientia Silvae Sinicae, 2012, 48 (7): 61- 65.
	王艳梅, 王海洋, 代莉, 等. 不同低温处理对12个种源山桐子种子休眠解除的影响. 山东农业大学学报: 自然科学版, 2015, 46 (1): 51- 56. doi: 10.3969/j.issn.1000-2324.2015.01.011
	Wang Y M , Wang H Y , Dai L , et al. Effect of different low temperature treatments on breaking Idesia polycarpa seed dormancy among 12 provenances. Journal of Shandong Agricultural University: Natural Science Edition, 2015, 46 (1): 51- 56. doi: 10.3969/j.issn.1000-2324.2015.01.011
	王艳梅, 姚兵, 刘伟伟, 等. 休眠解除后山桐子种子萌发过程中内源激素的动态变化. 林业科学, 2018, 54 (6): 44- 52.
	Wang Y M , Yao B , Liu W W , et al. Dynamic changes of endogenous hormones in seeds germination of Idesia polycarpa after dormancy release. Scientia Silvae Sinicae, 2018, 54 (6): 44- 52.
	魏俊, 陆秀君, 张晓林, 等. MicroRNA在种子发育, 休眠与萌发过程中的作用. 遗传, 2017a, 39 (1): 14- 21.
	Wei J , Lu X J , Zhang X L , et al. Functions of microRNA in seed development, dormancy and germination process. Hereditas, 2017, 39 (1): 14- 21.
	魏俊. 2017. 天女木兰种子萌发过程microRNA挖掘及其表达. 沈阳: 沈阳农业大学硕士学位论文.
	Wei J. 2017. Identification and expression analysis of microRNA during seed germination in Magnolia sieboldii K. Koch. Shenyang: MS thesis of Shenyang Agricultural University. [in Chinese]
	肖志成, 高捍东. 三角槭种子休眠与萌发特性研究. 西南林学院学报, 2008, 28 (5): 35- 38.
	Xiao Z C , Gao H D . Study on seed dormancy and germination characteristics of Acer buergerianum. Journal of Southwest Forestry College, 2008, 28 (5): 35- 38.
	许佳. 2016. 拟南芥miR393通过与ABA信号途径调控种子成熟和萌发的机制研究. 杭州: 浙江大学硕士学位论文.
	Xu J. 2016. MiR393 regulates seed maturation and germination through interaction with ABA in Arabidopsis thaliana. Hangzhou: MS thesis of Zhejiang University. [in Chinese]
	杨期和, 廖富林, 温献环, 等. 三药槟榔种子休眠与萌发的研究. 广西植物, 2005, 25 (6): 549- 554. doi: 10.3969/j.issn.1000-3142.2005.06.009
	Yang Q H , Liao F L , Wen X H , et al. Study on seed dormancy and germination of Areca catechu L. Guihaia, 2005, 25 (6): 549- 554. doi: 10.3969/j.issn.1000-3142.2005.06.009
	杨万霞. 2004. 青钱柳种子休眠原因及萌发生理的初步研究. 南京: 南京林业大学硕士学位论文.
	Yang W X. 2004. Preliminary study on seed dormancy and germination physiology of Cyclocarya paliurus. Nanjing: MS thesis of Nanjing Forestry University. [in Chinese]
	杨孝汉, 胡婉仪. 玉兰种子休眠解除过程中的生物大分子变化. 园艺学报, 1991, 18 (1): 75- 80. doi: 10.3321/j.issn:0513-353X.1991.01.007
	Yang X H , Hu W Y . Changes of biomacromolecules in Magnolia grandiflora seeds during dormancy breaking. Acta Horticulturae Sinica, 1991, 18 (1): 75- 80. doi: 10.3321/j.issn:0513-353X.1991.01.007
	杨艳华. 2009. 南京椴种子休眠解除过程胚乳物质代谢的研究. 南京: 南京林业大学硕士学位论文.
	Yang Y H. 2009. Studies on the relationship of substance metabolism and dormancy releasing of Tilia miqueliana seeds. Nanjing: MS thesis of Nanjing Forestry University. [in Chinese]
	尹华军, 刘庆. 种子休眠与萌发的分子生物学的研究进展. 植物学通报, 2004, 2, 156- 163. doi: 10.3969/j.issn.1674-3466.2004.02.004
	Yin H J , Liu Q . Advances in molecular biology of seed dormancy and germination. Chinese Bulletin of Botany, 2004, (2): 156- 163. doi: 10.3969/j.issn.1674-3466.2004.02.004
	游双红. 2013. 圆齿野鸦椿种子休眠机理及催芽技术研究. 南昌: 江西农业大学硕士学位论文.
	You S H. 2013. The dormancy mechanism and stratification technique of Euscaphis konishii seeds. Nanchang: MS thesis of Jiangxi Agricultural University. [in Chinese]
	于海莲. 2009. 南方红豆杉种子休眠机理及催芽技术的研究. 北京: 北京林业大学硕士学位论文.
	Yu H L. 2009. The seed dormancy mechanisms and germinating conditions of Taxus chinensis var. mairer. Beijing: MS thesis of Beijing Forestry University. [in Chinese]
	岳佳铭, 李曼莉. 萌发过程中种子贮藏物质的动员和变化规律概述. 种子, 2021, 40 (1): 56- 61.
	Yue J M , Li M L . Review of the mobilization and change of seed storage materials during germination. Seed, 2021, 40 (1): 56- 61.
	张存虎. 2018. 榉树种子打破休眠与萌发机理的研究. 泰安: 山东农业大学硕士学位论文.
	Zhang C H. 2018. The research of breaking dormancy and germination mechanism of Zelkova schneideriana seeds. Tai'an: MS thesis of Shandong Agricultural University. [in Chinese]
	张宁, 刘震. 冬季低温对耐冬山茶种子萌发的影响. 河南科学, 2015, 33 (1): 45- 48.
	Zhang N , Liu Z . Effect of low temperature in winter on the germination of seeds in Camellia japonica. Henan Science, 2015, 33 (1): 45- 48.
	赵东兴, 李春, 李芹, 等. H₂O₂和种皮处理对马槟榔种子萌发和幼苗生长的影响. 种子, 2018, 37 (8): 72- 74. doi: 10.3969/j.issn.1005-2690.2018.08.057
	Zhao D X , Li C , Li Q , et al. Effects of H₂O₂ soak and seed-coat treatment on seed germination and seedling growth of Capparis masaikai. Seed, 2018, 37 (8): 72- 74. doi: 10.3969/j.issn.1005-2690.2018.08.057
	郑光华. 几种园林植物种子休眠与发芽生理的研究. 园艺学报, 1963, 2 (3): 75- 88.
	Zheng G H . Study on seed dormancy and germination physiology of several garden plants. Acta Horticulturae Sinica, 1963, 2 (3): 75- 88.
	周峰. 燃烧植物产生的化学信号物质与种子萌发研究进展. 种子, 2013, 32 (9): 45- 47.
	Zhou F . Research advances in the relationship between seed germination and chemical signals from burning plants. Seed, 2013, 32 (9): 45- 47.
	周健. 2016. 紫荆种子休眠特性及其吸水机制研究. 南京: 南京林业大学硕士学位论文.
	Zhou J. 2016. The mechanism of water absorbing and dormancy in Cercis chinensis seeds. Nanjing: MS thesis of Nanjing Forestry University. [in Chinese]
	周学权. 油松种子萌发过程中核酸和蛋白质的变化. 北京林业大学学报, 1989, 11 (3): 46- 53.
	Zhou X Q . Changes of nucleic acid and protein during seed germination of Pinus tabulaeformis. Journal of Beijing Forestry University, 1989, 11 (3): 46- 53.
	Anand A , Kumari A , Thakur M , et al. Hydrogen peroxide signaling integrates with phytohormones during the germination of magnetoprimed tomato seeds. Scientific Reports, 2019, 9 (3): 1- 11.
	Barrero J M , Millar A A , Griffiths J , et al. Gene expression profiling identifies two regulatory genes controlling dormancy and ABA sensitivity in Arabidopsis seeds. The Plant Journal, 2010, 61 (4): 611- 622. doi: 10.1111/j.1365-313X.2009.04088.x
	Baskin J M , Baskin C C . A classification system for seed dormancy. Seed Science Research, 2004, 14 (1): 1- 16. doi: 10.1079/SSR2003150
	Bewley J D . Seed germination and dormancy. Plant Cell, 1997a, 9 (7): 1055- 1066.
	Bewley J D . Breaking down the walls-A role for endo-β-mannanase in release from seed dormancy?. Trends in Plant Science, 1997b, 2 (12): 464- 469. doi: 10.1016/S1360-1385(97)01147-3
	Bewley J D , Bradford K J , Hilhorst H W M , et al. Physiology of development, germination and dormancy 3rd edition. New York, 2013,
	Cao D C , Xu H M , Zhan Y Y , et al. Transcriptome and degradome sequencing reveals dormancy mechanisms of Cunninghamia lanceolata seeds. Plant Physiology, 2016, 172 (4): 347- 2362.
	Chapman E J , Estelle M . Mechanism of auxin-regulated gene expression in plants. Annu Review of Genetics, 2009, 43 (1): 265- 285. doi: 10.1146/annurev-genet-102108-134148
	Chen C , Twito S , Miller G . New cross talk between ROS, ABA and auxin controlling seed maturation and germination unraveled in APX6 deficient Arabidopsis seeds. Plant Signaling and Behavior, 2014, 9 (12): e976489. doi: 10.4161/15592324.2014.976489
	Chen L , Wang T Z , Zhao M G , et al. Ethylene-responsive miRNAs in roots of Medicago truncatula identified by high-throughput sequencing at whole genome level. Plant Science, 2012, 184, 14- 19. doi: 10.1016/j.plantsci.2011.11.007
	Chen L T , Wu K . Role of histone deacetylases HDA6 and HDA19 in ABA and abiotic stress response. Plant Signaling and Behavior, 2010, 5 (10): 1318- 1320. doi: 10.4161/psb.5.10.13168
	Chien C T , Chen S Y , Chang S H , et al. Dormancy and geimination in seeds of the medicinal Asian tree species Phellodendrort amurense var.wilsonii (Rutaceae). Seed Science and Technology, 2006, 34 (3): 561- 571. doi: 10.15258/sst.2006.34.3.03
	Chung P J , Park B S , Wang H , et al. Light-inducible MiR163 targets PXMT1 transcripts to promote seed germination and primary root elongation in Arabidopsis. Plant Physiology, 2016, 170 (3): 1772- 1782. doi: 10.1104/pp.15.01188
	Crocker W . Mechanics of dormancy in seeds. American Journal of Botany, 1916, 3 (3): 99- 120. doi: 10.1002/j.1537-2197.1916.tb05406.x
	Curaba J , Singh M B , Bhalla P L . MiRNAs in the crosstalk between phytohormone signalling pathways. Journal of Experimental Botany, 2014, 65 (6): 1425- 1438. doi: 10.1093/jxb/eru002
	Das S S , Karmakar P , Nandi A K , et al. Small RNA mediated regulation of seed germination. Frontiers in Plant Science, 2015, 6, 828.
	Davière J M , Achard P . A pivotal role of DELLAs in regulating multiple hormone signals. Molecular Plant, 2016, 9 (1): 10- 20. doi: 10.1016/j.molp.2015.09.011
	Eckstein L , Fenner M , Thompson K . The ecology of seeds. Basic and Applied Ecology, 2005, 6 (5): 488. doi: 10.1016/j.baae.2005.08.001
	Finkelstein R , Reeves W , Ariizumi T , et al. Molecular aspects of seed dormancy. Annual Review of Plant Biology, 2008, 59 (1): 387- 415. doi: 10.1146/annurev.arplant.59.032607.092740
	Flematti G R , Ghisalberti E L , Dixon K W , et al. A compound from smoke that promotes seed germination. Science, 2004, 305 (5686): 977. doi: 10.1126/science.1099944
	Footitt S , Douterelo-Soler I , Clay H , et al. Dormancy cycling in Arabidopsis seeds is controlled by seasonally distinct hormone-signaling pathways. Proceedings of the National Academy of Sciences, 2011, 108 (50): 20236- 20241. doi: 10.1073/pnas.1116325108
	Gonzalez G M , Abia D , Salinas J , et al. Two new alleles of the abscisic aldehyde oxidase 3 gene reveal its role in abscisic acid biosynthesis in seeds. Plant Physiology, 2004, 135 (1): 325- 333. doi: 10.1104/pp.103.036590
	Graeber K , Linkies A , Steinbrecher T , et al. "DELAY OF GERMINATION1" mediates a conserved coat-dormancy mechanism for the temperature- and gibberellin-dependent control of seed germination. Proceedings of the National Academy of Sciences, 2014, 111 (34): E3571- 3580. doi: 10.1073/pnas.1403851111
	Graeme E , Hobson . Changes in mitochondrial composition and behaviour in relation to dormancy. Annals of Applied Biology, 1981, 98 (3): 541- 544. doi: 10.1111/j.1744-7348.1981.tb00788.x
	Gu Y , Liu Y H , Zhang J J , et al. Identification and characterization of microRNAs in the developing maize endosperm. Genomics, 2013, 102 (5-6): 472- 478. doi: 10.1016/j.ygeno.2013.08.007
	Guan C M , Wang X C , Feng J , et al. Cytokinin antagonizes abscisic acid mediated inhibition of cotyledon greening by promoting the degradation of abscisic acid insensitive 5 protein in Arabidopsis. Plant Physiology, 2014, 164 (3): 1515- 1526. doi: 10.1104/pp.113.234740
	Harper J L . The ecological significance of dormancy and its importance in weed control. Proceedings of the IVth International Congress of Crop Protection, 1957, 35 (1): 415- 420.
	He J , Chen Q , Xin P , et al. Author Correction: CYP72A enzymes catalyse 13-hydrolyzation of gibberellins. Nature Plants, 2019, 5 (10): 69.
	Hu Y , Yu D . Brassinosteroid Insensitive2 interacts with Abscisic Acid Insensitive5 to mediate the antagonism of brassinosteroids to abscisic acid during seed germination in Arabidopsis. The Plant Cell, 2014, 26 (11): 4394- 4408. doi: 10.1105/tpc.114.130849
	Huang Y , Feng C Z , Ye Q , et al. Arabidopsis WRKY6 transcription factor acts as a positive regulator of abscisic acid signaling during seed germination and early seedling development. PLoS Genetics, 2016, 12 (2): e1005833. doi: 10.1371/journal.pgen.1005833
	Huang D Q , Koh C , Feurtado J A , et al. MicroRNAs and their putative targets in Brassica napus seed maturation. BMC Genomics, 2013, 14 (1): 1- 25. doi: 10.1186/1471-2164-14-1
	Jacobsen J V , Barrero J M , Hughes T , et al. Roles for blue light, jasmonate and nitric oxide in the regulation of dormancy and germination in wheat grain (Triticum aestivum L). . Planta, 2013, 238 (1): 121- 138. doi: 10.1007/s00425-013-1878-0
	Jia Z , Zhao B , Liu S , et al. Embryo transcriptome and miRNA analyses reveal the regulatory network of seed dormancy in Ginkgo biloba. Tree Physiology, 2020, 41 (4)
	Jian Z , Chen F , Zhi W , et al. A novel role for histone methyltransferase KYP/SUVH4 in the control of Arabidopsis primary seed dormancy. New Phytologist, 2012, 193 (3): 605- 616. doi: 10.1111/j.1469-8137.2011.03969.x
	Ju L , Jing Y X , Shi P T , et al. JAZ proteins modulate seed germination through interaction with ABI5 in breadwheat and Arabidopsis. New Phytologist, 2019, 223 (1): 246- 260. doi: 10.1111/nph.15757
	Keeley J E , Fotheringham C J . Trace gas emissions and smoke-induced seed germination. Science, 1997, 276 (5316): 1248- 1250. doi: 10.1126/science.276.5316.1248
	Kim W , Lee Y , Park J , et al. HONSU, a protein phosphatase 2C, regulates seed dormancy by inhibiting ABA signaling in Arabidopsis. Plant and Cell Physiology, 2013, 54 (4): 555- 572. doi: 10.1093/pcp/pct017
	Lee S , Cheng H , King K E , et al. Gibberellin regulates Arabidopsis seed germination via RGL2, a GAI/RGA-like gene whose expression is up-regulated following gimbibition. Genes and Development, 2002, 16 (5): 646- 658. doi: 10.1101/gad.969002
	Lee S , Kim S G , Park C M . Salicylic acid promotes seed germination under high salinity by modulating antioxidant activity in Arabidopsis. New Phytologist, 2010, 188 (2): 626- 637. doi: 10.1111/j.1469-8137.2010.03378.x
	Lee K , Lee H G , Yoon S , et al. The ArabidopsisMYB96 transcription factor is a positive regulator of Abscisic Acid-Insensitive 4 in the control of seed germination. Plant Physiology, 2015a, 168 (2): 677- 689. doi: 10.1104/pp.15.00162
	Lee S J , Lee M H , Kim J I , et al. Arabidopsis putative MAP kinase kinase kinases RAF10 and RAF11 are positive regulators of seed dormancy and ABA response. Plant and Cell Physiology, 2015b, 56 (1): 84- 97. doi: 10.1093/pcp/pcu148
	Li B L , Foley M E . Genetic and molecular control of seed dormancy. Trends in Plant Science, 1997, 2 (10): 384- 389. doi: 10.1016/S1360-1385(97)90053-4
	Li D , Liu Z , Gao L , et al. Genome-wide identification and characterization of micrornas in developing grains of Zea mays L. PLoS ONE, 2016, 11 (4): e0153168. doi: 10.1371/journal.pone.0153168
	Li Z , Gao Y , Zhang Y C , et al. Reactive oxygen species and gibberellin acid mutual induction to regulate tobacco seed germination. Frontiers in Plant Science, 2018, 9, 1- 15. doi: 10.3389/fpls.2018.00001
	Li Z , Xu J G , Gao Y , et al. The synergistic priming effect of exogenous salicylic acid and H₂O₂ on chilling tolerance enhancement during maize (Zea mays L. ) seed germination. Frontiers in Plant Science, 2017, 8, 1153. doi: 10.3389/fpls.2017.01153
	Li X Y , Chen T T , Li Y , et al. ETR1/RDO3 regulates seed dormancy by relieving the inhibitory effect of the ERF12-TPL complex on DELAY OF GERMINATION1 expression. The Plant Cell, 2019, 31, 832- 847. doi: 10.1105/tpc.18.00449
	Linkies A , Muller K , Morris K , et al. Ethylene interacts with abscisic acid to regulate endosperm rupture during germination: a comparative approach using Lepidium sativum and Arabidopsis thaliana. The Plant Cell, 2009, 21 (12): 3803- 3822.
	Liu H , Guo S , Lu M , et al. Biosynthesis of DHGA12 and its roles in Arabidopsis seedling establishment. Nature communications, 2019, 10, 1- 14. doi: 10.1038/s41467-018-07882-8
	Liu X D , Zhang H , Zhao Y , et al. Auxin controls seed dormancy through stimulation of abscisic acid signaling by inducing ARF-mediated ABI3 activation in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America, 2013a, 110 (38): 15485- 15490. doi: 10.1073/pnas.1304651110
	Liu X , Hu P , Huang M , et al. The NF-YC-RGL2 module integrates GA and ABA signaling to regulate seed germination in Arabidopsis. Nature Communications, 2016, 7, 1- 14.
	Liu Y , Koornneef M , Soppe W . The absence of histone H₂B monoubiquitination in the Arabidopsis Hub1 (rdo4) mutant reveals a role for chromatin remodeling in seed dormancy. The Plant Cell, 2007, 19, 433- 444. doi: 10.1105/tpc.106.049221
	Liu Y J , Xiu Z H , Meeley R , et al. Empty Pericarp5 encodes a pentatricopeptide repeat protein that is required for mitochondrial RNA editing and seed development in maize. Plant Cell, 2013, 25 (3): 868- 883. doi: 10.1105/tpc.112.106781
	Long J A , Ohno C , Smith Z R , et al. Topless regulates apical embryonic fate in Arabidopsis. Science, 2006, 312, 1520- 1523. doi: 10.1126/science.1123841
	Ma Y , Szostkiewicz I , Korte A , et al. Regulators of PP2C phosphatase activity function as abscisic acid sensors. Science, 2009, 324, 1064- 1068. doi: 10.1126/science.1172408
	Magome H , Yamaguchi S , Hanada A , et al. The DDF1 transcriptional activator upregulates expression of a gibberellin-deactivating gene, GA2ox7, under high-salinity stress in Arabidopsis. Plant Journal, 2008, 56, 613- 626. doi: 10.1111/j.1365-313X.2008.03627.x
	Nazir , Hussain , Naseer , et al. Maturity indices and dormancy breaking methods of black locust (Rhobinia pseudoacacia) seeds under temperate Kashmir condition. Ecology Environment and Conservation, 2014, 20 (4): 1769- 1775.
	Née G , Kramer K , Nakabayashi K. , et al. DELAY OF GERMINATION1 requires PP2C phosphatases of the ABA signalling pathway to control seed dormancy. Nature Communication, 2017, 8 (1): 72. doi: 10.1038/s41467-017-00113-6
	Nikolaeva M G . Factors controlling the seed dormancy pattern. Physiology and Biochemistry of Seed Dormancy and Germination, 1977, 51- 74.
	Nonogaki H . Seed dormancy and germination-emerging mechanisms and new hypotheses. Front in Plant Science, 2014, 5, 233.
	Nonogaki H . MicroRNA gene regulation cascades during early stages of plant development. Plant Cell Physiology, 2010, 51 (11): 1840- 1846. doi: 10.1093/pcp/pcq154
	Noriyuki N , Wataru T , Moresco J J , et al. Control of seed dormancy and germination by DOG1-AHG1 PP2C phosphatase complex via binding to heme. Nature Communications, 2018, 9 (1): 2132. doi: 10.1038/s41467-018-04437-9
	Pan J , Wang H , Hu Y , et al. ArabidopsisVQ18 and VQ26 proteins interact with ABI5 transcription factor to negatively modulate ABA response during seed germination. The Plant Journal, 2018, 95 (3): 529- 544. doi: 10.1111/tpj.13969
	Park J , Kim Y S , Kim S G , et al. Integration of auxin and salt signals by the NAC transcription factor NTM2 during seed germination in Arabidopsis. Plant Physiology, 2011, 156 (2): 537- 549. doi: 10.1104/pp.111.177071
	Pipinis E , Stampoulidis A , Milios E , et al. Effects of seed moisture content, stratification and sowing date on the germination of Corylus avellana seeds. Journal of Forestry Research, 2020, 31 (3): 743- 749. doi: 10.1007/s11676-018-0852-x
	Piskurewicz U , Jikumaru Y , Kinoshita N , et al. The gibberellic acid signaling repressor RGL2 inhibits Arabidopsis seed germination by stimulating abscisic acid synthesis and ABI5 activity. Plant Cell, 2008, 20 (10): 2729- 2745. doi: 10.1105/tpc.108.061515
	Qi J S , Song C P , Wang B , et al. Reactive oxygen species signaling and stomatal movement in plant responses to drought stress and pathogen attack. Journal of Integrative Plant Biology, 2018, 60 (9): 805- 826. doi: 10.1111/jipb.12654
	Qi Y , Bilan M V , Chin K L . New method for breaking Korean pine seed dormancy. Journal of Arboriculture, 1993, 19 (2): 113- 117.
	Remington D L , Vision T J , Guilfoyle T J , et al. Contrasting modes of diversification in the Aux/IAA and ARF gene families. Plant Physiology, 2004, 135 (3): 1738- 1752. doi: 10.1104/pp.104.039669
	Resentini F , Felipo-Benavent A , Colombo L , et al. TCP14 and TCP15 mediate the promotion of seed germination by gibberellins in Arabidopsis thaliana . Molecular Plant, 2015, 8 (3): 482- 485. doi: 10.1016/j.molp.2014.11.018
	Reyes J L , Chua N H . ABA induction of MIR159 controls transcript levels of two MYB factors during Arabidopsis seed germination. Plant journal, 2007, 49 (4): 592- 606. doi: 10.1111/j.1365-313X.2006.02980.x
	Roberts E H , Smith R D . Viability of seeds. London: Chapman and Hall Ltd, 1977,
	Rui Z , Sun H L , Mei C , et al. The Arabidopsis Ca²⁺-dependent protein kinase CPK12 negatively regulates abscisic acid signaling in seed germination and post-germination growth. New Phytologist, 2011, 192 (1): 61- 73. doi: 10.1111/j.1469-8137.2011.03793.x
	Russell L , Larner V , Kurup S , et al. The Arabidopsis comatose locus regulates germination potential. Development, 2000, 127 (17): 3759- 3767. doi: 10.1242/dev.127.17.3759
	Shu K , Zhang H W , Wang S F , et al. ABI4 regulates primary seed dormancy by regulating the biogenesis of abscisic acid and gibberellins in Arabidopsis. PLoS Genetics, 2013, 9 (6): e1003577. doi: 10.1371/journal.pgen.1003577
	Thole J M , Beisner E R , Liu J , et al. Abscisic acid regulates root elongation through the activities of auxin and ethylene in Arabidopsis thaliana. G3 (Genes Genomes Genetics), 2014, 4 (7): 1259- 1274.
	Valérie L , North H , Frey A , et al. Functional analysis of Arabidopsis NCED6 and NCED9 genes indicates that ABA synthesized in the endosperm is involved in the induction of seed dormancy. Plant Journal for Cell and Molecular Biology, 2010, 45 (3): 309- 319.
	Villiers T A . Dormancy and the survival of plants. Edward Arnold, 1975, 30 (2): 67.
	Vleeshouwers L , Bouwmeester H , Karssen C . Redefining seed dormancy: an attempt to integrate physiology and ecology. Journal of Ecology, 1995, 83 (6): 1031- 1037. doi: 10.2307/2261184
	Wang L , Hua D P , He J N , et al. Auxin Response Factor2 (ARF2) and its regulated homeodomain gene HB33 mediate abscisic acid response in Arabidopsis. PLoS Genetics, 2011a, 7 (7): e1002172. doi: 10.1371/journal.pgen.1002172
	Wang Y M , Liu W W , Wang D H , et al. Study on the effect of different photoperiods on the seedling growth of Idesia policarpa Maxim. from different provenances. Fresenius Environmental Bulletin, 2019a, 28 (2A): 1229- 1234.
	Wang Y M , Liu Z , Li F . The dynamic changes of plant endogenous hormones in response to different temperatures in seeds of Idesia polycarpa Maxim. from Xinyang. Fresenius Environmental Bulletin, 2019b, 28 (4): 2495- 2505.
	Wang Y M , Wang L J , Yao B , et al. Changes in ABA, IAA, GA₃, and ZR levels during seed dormancy release in Idesia polycarpa Maxim. from Jiyuan. Polish Journal of Environmental Studies, 2018, 27 (4): 1833- 1839. doi: 10.15244/pjoes/78041
	Wang Y P , Li L , Ye T T , et al. Cytokinin antagonizes ABA suppression to seed germination of Arabidopsis by down regulating ABI5 expression. The Plant Journal, 2011b, 68 (2): 249- 261. doi: 10.1111/j.1365-313X.2011.04683.x
	Wang Z , Cao H , Sun Y , et al. Arabidopsis paired amphipathic helix proteins SNL1 and SNL2 redundantly regulate primary seed dormancy via abscisic acid-ethylene antagonism mediated by histone deacetylation. The Plant Cell, 2013, 25 (1): 149- 166. doi: 10.1105/tpc.112.108191
	Wareing P F , Saunders P F . Hormones and dormancy. Annual Review of Plant Physiology, 1971, 22 (1): 261- 288. doi: 10.1146/annurev.pp.22.060171.001401
	Xie Z , Zhang Z L , Hanzlik S , et al. Salicylic acid inhibits gibberellin-induced alpha-amylase expression and seed germination via a pathway involving an abscisic-acid-inducible WRKY gene. Plant Molecular Biology, 2007, 64 (3): 293- 303. doi: 10.1007/s11103-007-9152-0
	Xie Z X , Allen E , Fahlgren N , et al. Expressionof Arabidopsis MiRNA genes. Plant Physiology, 2005, 138, 2145- 2154. doi: 10.1104/pp.105.062943
	Xing L , Zhu M , Zhang M , et al. High-throughput sequencing of small RNA transcriptomes in maize kernel identifies miRNAs involved in embryo and endosperm development. Genes, 2017, 8 (12): 385. doi: 10.3390/genes8120385
	Yan A , Chen Z . The control of seed dormancy and germination by temperature, light and nitrate. The Botanical Review, 2020, 86 (1): 39- 75. doi: 10.1007/s12229-020-09220-4
	Zabala G , Campos E , Varala K K , et al. Divergent patterns of endogenous small RNA populations from seed and vegetative tissues of Glycine max. BMC Plant Biology, 2012, 12 (1): 177. doi: 10.1186/1471-2229-12-177
	Zhang J H , Zhang S G , Han S Y , et al. Deciphering small noncoding RNAs during the transition from dormant embryo to germinated embryo in Larches (Larix leptolepis). PLoS ONE, 2013, 8 (12): e81452. doi: 10.1371/journal.pone.0081452
	Zhao M , Yang S , Liu X , et al. Arabidopsis histone demethylases LDL1 and LDL2 control primary seed dormancy by regulating DELAY of GERMINATION1 and ABA signaling-related genes. Frontiers in Plant Science, 2015, 6, 159.
	Zhou C , Lin Q , Lan J , et al. WRKY transcription factor OsWRKY29 represses seed dormancy in rice by weakening abscisic acid response. Frontiers in Plant Science, 2020, 11, 691. doi: 10.3389/fpls.2020.00691
	Zhu M , Zhang M , Xing L , et al. Transcriptomic analysis of long non-coding RNAs and coding genes uncovers a complex regulatory network that is involved in Maize seed development. Genes, 2017, 8 (10): 274. doi: 10.3390/genes8100274
	Zhu S Y , Yu X C , Wang X J , et al. Two calcium-dependent protein kinases, CPK4 and CPK11, regulate abscisic acid signal transduction in Arabidopsis. The Plant Cell, 2007, 19, 3019- 3036. doi: 10.1105/tpc.107.050666

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

[1]	Chen Chen,Fangyuan Yu. Research Progress on Flower Bud Differentiation of Trees [J]. Scientia Silvae Sinicae, 2020, 56(9): 119-129.
[2]	Dejie Yin,Fengqin Bu,Yanfang Xu,Biao Huang,Cheng Li. Dormancy Characteristics and Breaking Methods of Vitex rotundifolia Seeds [J]. Scientia Silvae Sinicae, 2020, 56(12): 157-165.
[3]	Yang Yang,Haiyang Wang,Lihui Ma. Characteristics of Fruiting and Seed Germination of Endangered Plant, Rhododendron changii [J]. Scientia Silvae Sinicae, 2020, 56(10): 173-183.
[4]	Jiang Zheng, Yu Qiannan, Qiao Mingfeng, Xiao Juan, Zhang Ziliang, Yin Huajun. Effects of Root Exudates from Picea asperata Seedlings on the Seed Germination and Seedling Growth of Two Herb Species [J]. Scientia Silvae Sinicae, 2019, 55(6): 160-166.
[5]	Liangang Zhang,Ling Zhi,Guangyu Wang,Ya Li. Influencing Factors and Improving Path of Farmers' Satisfaction with the Grain for Green Project: A Case Study of Heqing County in Yunnan Province and Zhijin County in Guizhou Province [J]. Scientia Silvae Sinicae, 2019, 55(12): 123-132.
[6]	Zheng Jing, Lin Huiqi. Forest Farmers’ Cognition and Its Influence Factors of Commercial Forest Redemption in Key Ecological Location ——Based on Case Study from Fujian Province [J]. Scientia Silvae Sinicae, 2018, 54(9): 114-124.
[7]	Luo Xiaoman, Ding Guijie, Wang Yi. Active Constituents of Soil Extracts from Mycorrhizal Seedling Rhizosphere of Pinus massoniana and Their Effects on Seed Germination [J]. Scientia Silvae Sinicae, 2018, 54(8): 32-38.
[8]	Wang Yanmei, Yao Bing, Liu Weiwei, Wang Lijun, Yan Huiping, Li Fei, Liu Zhen. Dynamic Changes of Endogenous Hormones in Seeds Germination of Idesia polycarpa after Dormancy Release [J]. Scientia Silvae Sinicae, 2018, 54(6): 44-52.
[9]	Zhang Junjie, Chai Shengfeng, Wei Xiao, Lv Shihong, Wu Shaohua. Germination Characteristics of the Seed of a Rare and Endangered Plant, Garcinia paucinervis [J]. Scientia Silvae Sinicae, 2018, 54(4): 174-185.
[10]	Zhang Peng, Zhao Tongtong, Li Mingyue, Wu Lingdong, Shen Hailong. Effects of CaCl₂, ABA or Sucrose Pretreatment on Germination of Dormancy Released Seeds of Fraxinus mandshurica Dehydrated at Low Temperature [J]. Scientia Silvae Sinicae, 2017, 53(7): 54-61.
[11]	Chen Xiangbo, Liu Yang, Zhao Mingshui, Tu Shuping. Embryo Development and Dormancy Releasing of Acer yangjuechi, the Extremely Endangered Plant [J]. Scientia Silvae Sinicae, 2017, 53(4): 65-73.
[12]	Zhang Peng, Song Boyang, Wu Lingdong, Shen Hailong. Germination and Physiological Responses of Dormancy Released Seeds of Fraxinus mandshurica to Dehydration in Different Conditions [J]. Scientia Silvae Sinicae, 2017, 53(3): 60-67.
[13]	Yan Fang, Zhang Enhe, Wang Qinli, Mao Zhuhong. The Mechanisms of Seed Dormancy of Wild Daphne giraldii and Methods for Dormancy Breaking [J]. Scientia Silvae Sinicae, 2016, 52(4): 30-37.
[14]	Liao Wenmei, Zhang Guanglai, Kong Fanbin. Farmers' Demands on Forestry Socialized Services and its Influencing Factors—A Survey Based on 1 413 Households in 8 Provinces [J]. Scientia Silvae Sinicae, 2016, 52(11): 148-156.
[15]	Du Yuliang, Liu Caihong, Chen Ye, Yin Zengfang. Establishment and Influencing Factors of in vitro Culture System for Tracheary Elements Differentiation [J]. Scientia Silvae Sinicae, 2014, 50(8): 146-153.

Research Progress on Dormancy and Germination Mechanism of Forest Seeds

RichHTML

PDF (PC)

Knowledge

Abstract

Cite this article

share this article

Figures/Tables 7

References 0

Related Articles 15

Recommended Articles

Metrics

Comments