Progress in research on the physiological and molecular mechanisms of seed dormancy in legumes

doi:10.11686/cyxb2025337

Abstract

Abstract:

Seed dormancy is an adaptive trait that developed during long-term evolution， enabling plants to survive under adverse conditions. Legumes are a core crop group in global agricultural production （encompassing food， forage， and green manure）， and the dormancy characteristics of their seeds directly affect sowing quality， field emergence rate， and yield stability. The molecular regulation of seed dormancy in legumes primarily revolves around the balance between abscisic acid （ABA） and gibberellin （GA） levels， involving a complex system that integrates the expression of genes involved in seed coat development and transcription factor networks， as well as epigenetic modifications and environmental signals. This mechanism ensures seed survival under unfavorable conditions while enabling precise germination under suitable circumstances. Physiological dormancy is the primary factor in seed dormancy， where ABA and GA do not directly interact but instead exhibit antagonistic effects-ABA promotes “dormancy maintenance” while GA triggers “germination initiation”. In Arabidopsis thaliana， many downstream targets of the ABA-responsive germination inhibitor ABI3 have been identified， including the ABI5， DELLA， and bHLH transcription factors. However， the corresponding regulatory network in legumes remains unclear， and further research is required to explore the interactions between ABI3 and other factors in the ABA signaling pathway. Based on recent research， this paper systematically reviews the types of seed dormancy in legumes， dormancy-breaking techniques， and molecular regulatory networks， with a focus on legume-specific dormancy traits. Furthermore， drawing on advances in research on the molecular regulation of seed dormancy in model plants， we propose a hypothetical model for the molecular regulation of seed dormancy in legumes. This review provides a theoretical foundation for understanding the molecular mechanisms of seed dormancy in legumes while offering insights for cultivation practices and varietal improvement.

Key words: legume seeds, physical dormancy, physiological dormancy, seed dormancy release, molecular regulatory mechanisms

Hao-zhen LIU, Shi-qin ZHAO, Shu-rong FENG, Cheng WANG, Jing-hong ZHANG, Shou-jiang SUN. Progress in research on the physiological and molecular mechanisms of seed dormancy in legumes[J]. Acta Prataculturae Sinica, 2026, 35(8): 207-220.

Figures/Tables 6

References 109

[1]	Hilhorst H W M. A critical update on seed dormancy. I. Primary dormancy. Seed Science Research， 1995， 5（2）： 61-73.
[2]	Smykal P， Vernoud V， Blair M W， et al. The role of the testa during development and in establishment of dormancy of the legume seed. Frontiers in Plant Science， 2014， 5： 351.
[3]	Savage W E F， Metzger G L. Seed dormancy and the control of germination. New Phytologist， 2006， 171（3）： 501-523.
[4]	Yang L W， Liu S R， Lin R C. The role of light in regulating seed dormancy and germination. Journal of Integrative Plant Biology， 2020， 62（9）： 1310-1326.
[5]	Huang X Q， Schmitt J， Dorn L， et al. The earliest stages of adaptation in an experimental plant population： Strong selection on QTLS for seed dormancy. Molecular Ecology， 2010， 19（7）： 1335-1351.
[6]	Donohue K， de Casas R R， Burghardt L， et al. Germination， postgermination adaptation， and species ecological ranges. Annual Review of Ecology， Evolution， and Systematics， 2010， 41（1）： 293-319.
[7]	Gary J M， Catherine F. Genetics and Genomics of the Triticeae. New York： Springer US， 2009： 31-79.
[8]	Gubler F， Millar A A， Jacobsen J V. Dormancy release， ABA and pre-harvest sprouting. Current Opinion in Plant Biology， 2005， 8（2）： 183-187.
[9]	Bewley J D. Breaking down the walls-a role for endo-β-mannanase in release from seed dormancy？ Trends in Plant Science， 1997， 2（12）： 464-469.
[10]	Nonogaki H， Barrero J M， Li C. Editorial： Seed dormancy， germination， and pre-harvest sprouting. Frontiers in Plant Science， 2018， 9： 1783.
[11]	Malloch A J C， Bewley J D， Black M. Seeds： Physiology of development and germination. The Journal of Ecology， 1986， 74（3）： 905.
[12]	Vleeshouwers L M， Bouwmeester H J， Karssen C M. Redefining seed dormancy： An attempt to integrate physiology and ecology. The Journal of Ecology， 1995， 83（6）： 1031.
[13]	Batlla D， Benech-Arnold R L. Predicting changes in dormancy level in weed seed soil banks： Implications for weed management. Crop Protection， 2007， 26（3）： 189-197.
[14]	Nonogaki H. Seed dormancy and germination-emerging mechanisms and new hypotheses. Frontiers in Plant Science， 2014， 5： 233.
[15]	Smith S M， Weller J L. Seasonal control of seed germination. New Phytologist， 2019， 225（5）： 1821-1823.
[16]	Baskin J M， Baskin C C. A classification system for seed dormancy. Seed Science Research， 2004， 14（1）： 1-16.
[17]	Nautiyal P C， Sivasubramaniam K， Dadlani M. Seed dormancy and regulation of germination. Seed Science and Technology， 2023， 52： 39-66.
[18]	Baskin J M， Baskin C C. The great diversity in kinds of seed dormancy： A revision of the nikolaeva-baskin classification system for primary seed dormancy. Seed Science Research， 2021， 31（4）： 249-277.
[19]	Schellenberg M P. Seeds： Ecology， biogeography， and evolution of dormancy and germination. The Canadian Field-Naturalist， 1999， 113（4）： 702-703.
[20]	Amen R D. A model of seed dormancy. The Botanical Review， 1968， 34（1）： 1-31.
[21]	Wareing P F， Saunders P F. Hormones and dormancy. Annual Review of Plant Physiology， 1971， 22（1）： 261-288.
[22]	Hilhorst H W M， Finch-Savage W E， Buitink J， et al. Dormancy in plant seeds//Dormancy and resistance in harsh environments. Berilin： Springer， 2010： 43-67.
[23]	Cheng X J， Zhang S， E Z G， et al. Maternally expressed FERTILIZATION-INDEPENDENT ENDOSPERM1 regulates seed dormancy and aleurone development in rice. The Plant Cell， 2024， 37（1）： koae304.
[24]	Nautiyal P C， Bandyopadhyay A， Zala P V. In situ sprouting and regulation of fresh-seed dormancy in Spanish type groundnut （Arachis hypogaea L.）. Field Crops Research， 2001， 70（3）： 233-241.
[25]	Fennimore S. Genetic and physiological evidence for the role of gibberellic acid in the germination of dormant Avena fatua seeds. Journal of Experimental Botany， 1998， 49（318）： 89-94.
[26]	Kepczynski J， Kepczynska E. Ethylene in seed dormancy and germination. Physiologia Plantarum， 1997， 101（4）： 720-726.
[27]	Matilla A J. Ethylene in seed formation and germination. Seed Science Research， 2000， 10（2）： 111-126.
[28]	Li B， Foley M E. Differential polypeptide patterns in imbibed dormant and after-ripened Avena fatua embryos. Journal of Experimental Botany， 1994， 45（2）： 275-279.
[29]	Holdsworth M， Kurup S， Mkibbin R. Molecular and genetic mechanisms regulating the transition from embryo development to germination. Trends in Plant Science， 1999， 4（7）： 275-280.
[30]	Jayasuriya K M G G， Baskin J M， Geneve R L， et al. Sensitivity cycling and mechanism of physical dormancy break in seeds of Ipomoea hederacea （Convolvulaceae）. International Journal of Plant Sciences， 2009， 170（4）： 429-443.
[31]	Baskin J M， Baskin C C， Li X. Taxonomy， anatomy and evolution of physical dormancy in seeds. Plant Species Biology， 2000， 15（2）： 139-152.
[32]	Xue T T， Shen Y B， Liu J， et al. Advances in research of seed physical dormancy. Chinese Bulletin of Botany， 2016， 51（6）： 863-871.
	薛婷婷，沈永宝，刘嘉，等. 种子物理休眠研究进展. 植物学报， 2016， 51（6）： 863-871.
[33]	Zhang Y M， Ding X P， Zhang P， et al. Study on breaking dormancy in Astragalus sinicus L. Horticulture & Seed， 2011（5）： 19-20.
	张玉梅，丁显萍，张萍，等. 紫云英种子休眠破除方法的研究. 园艺与种苗， 2011（5）： 19-20.
[34]	Hu X W. Study on the mechanism of seed dormancy formation and breaking in leguminous plants.Lanzhou： Lanzhou University， 2008.
	胡小文. 豆科植物种子休眠形成与破除机制研究. 兰州：兰州大学， 2008.
[35]	Sedláková V， Hanáček P， Grulichová M， et al. Evaluation of seed dormancy， one of the key domestication traits in chickpea. Agronomy， 2021， 11（11）： 2292.
[36]	Kondo T， Sato C， Baskin J M， et al. Post-dispersal embryo development， germination phenology， and seed dormancy in Cardiocrinum cordatum var. glehnii （liliaceae s. str.）， a perennial herb of the broadleaved deciduous forest in Japan. American Journal of Botany， 2006， 93（6）： 849-859.
[37]	Sohindji F S， Sogbohossou D E O， Zohoungbogbo H P F， et al. Understanding molecular mechanisms of seed dormancy for improved germination in traditional leafy vegetables： An overview. Agronomy， 2020， 10（1）： 57.
[38]	Wang X M， Zhang H， Song R， et al. Multiple omics datasets reveal significant physical and physiological dormancy in alfalfa hard seeds identified by multispectral imaging analysis. The Crop Journal， 2023， 11（5）： 1458-1468.
[39]	Zhang X Y， Zou H， Zhang X X. Overview of research on causes of seed dormancy and methods of breaking. Biology Teaching， 2024， 49（9）： 2-5.
	张西园，邹慧，张欣欣. 种子休眠原因与破除研究概述. 生物学教学， 2024， 49（9）： 2-5.
[40]	Guo L P. Study on dormancy and dormancy breaking of tree peony seeds. Yangling： Northwest A&F University， 2016.
	郭丽萍. 种子休眠原因及休眠解除方法研究. 杨凌：西北农林科技大学， 2016.
[41]	Cui L L， Liu J Y， Wang Z L， et al. Study on the effects of different treatments on breaking the hard-seededness of new Medicago falcata variety. Seed， 2020， 39（6）： 24-30.
	崔乐乐，刘佳月，王照兰，等. 不同方法破除黄花苜蓿新品系种子硬实的效果研究.种子， 2020， 39（6）： 24-30.
[42]	Feng Y. Seed physical dormancy release in different populations of Sophora alopecuroides L. Lanzhou： Lanzhou University， 2011.
	冯燕. 苦豆子（Sophora alopecuroides L.）不同种群种子物理休眠释放的研究. 兰州：兰州大学， 2011.
[43]	Thanos C A， Georghiou K， Kadis C， et al. Cistaceae： A plant family with hard seeds. Israel Journal of Plant Sciences， 1992，41（4/5/6）： 251-263.
[44]	Zhang Y， Ma Y M， Qin J Y， et al. The study on seed dormancy and germination characteristics of Iris scariosa. Acta Agrestia Sinica， 2016， 24（5）： 1062-1067.
	张云，马玉梅，秦景逸，等. 膜苞鸢尾种子休眠及萌发特性研究. 草地学报， 2016， 24（5）： 1062-1067.
[45]	Fan P H. Research for seed dormancy breaking and allelopathy of Oxytropis ochrocephala Bunge. Xi’an： Northwest University，2014.
	樊鹏辉. 黄花棘豆的种子休眠破除及化感作用研究. 西安：西北大学， 2014.
[46]	Wang Y R， Hanson J. An improved method for breaking dormancy in seeds of Sesbania sesban. Experimental Agriculture， 2008， 44（2）： 185-195.
[47]	Auld T D， O'connell M A. Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae. Australian Journal of Ecology， 1991， 16（1）： 53-70.
[48]	Yu L， Wang Y R， Song J H. Effects of temperatures and pre-treatments on germination of Sorghum sudanense. Pratacultural Science， 1991（5）： 65-68.
	余玲，王彦荣，孙建华. 温度及预处理对苏丹草种子萌发的影响. 草业科学， 1991（5）： 65-68.
[49]	Li X， Baskin J M， Baskin C C. Anatomy of two mechanisms of breaking physical dormancy by experimental treatments in seeds of two north American Rhus species （Anacardiaceae）. American Journal of Botany， 1999， 86（11）： 1505-1511.
[50]	Hu X W， Wu Y P， Wang Y R， et al. Primary study of release method for legume seed dormancy. Acta Botanica Boreali-Occidentalia Sinica， 2009， 29（3）： 568-573.
	胡小文，武艳培，王彦荣，等. 豆科种子休眠破除方法初探. 西北植物学报， 2009， 29（3）： 568-573.
[51]	Wang J， Wang Z J， Chen Y， et al. The study on seed germination characteristic of Sophora alopecuroides. Agricultural Research in the Arid Areas， 2007， 25（4）： 202-206.
	王进，王泽基，陈叶，等. 苦豆子种子发芽特性研究. 干旱地区农业研究， 2007， 25（4）： 202-206.
[52]	Ghaghelestany A B， Alebrahim M T， MacGregor D R， et al. Evaluation of ultrasound technology to break seed dormancy of common lambsquarters （Chenopodium album）. Food Science & Nutrition， 2020， 8（6）： 2662-2669.
[53]	Peng C Y， Wu Y， Shi F H， et al. Review of the current research progress of seed germination inhibitors. Horticulturae， 2023， 9（4）： 462.
[54]	Wang S S， Shi F L， Shi R R， et al. Seed dormancy and germination in alfalfa （Medicago falcata L.）. Legume Research， 2024， 47（2）： 234-241.
[55]	Finch-Savage W E， Footitt S. Seed dormancy cycling and the regulation of dormancy mechanisms to time germination in variable field environments. Journal of Experimental Botany， 2017， 68（4）： 843-856.
[56]	Finkelstein R， Reeves W， Ariizumi T， et al. Molecular aspects of seed dormancy. Annual Review of Plant Biology， 2008， 59（1）： 387-415.
[57]	Soltani A， Walter K A， Wiersma A T， et al. The genetics and physiology of seed dormancy， a crucial trait in common bean domestication. BMC Plant Biology， 2021， 21（1）： 58.
[58]	García A G， Balarynová J， Smykal P， et al. Genetic and transcriptomic analysis of lentil seed imbibition and dormancy in relation to its domestication. The Plant Genome， 2025， 18（2）： e70021.
[59]	Forbes I， Wells H D. Hard and soft seededness in blue lupine， Lupinus angustifolius L.： Inheritance and phenotype classification. Crop Science， 1968， 8（2）： 195-197.
[60]	Kongjaimun A， Kaga A， Tomooka N， et al. The genetics of domestication of yardlong bean， Vigna unguiculata （L.） Walp. ssp. unguiculata cv.gr. sesquipedalis. Annals of Botany， 2012， 109（6）： 1185-1200.
[61]	Weeden N F. Genetic changes accompanying the domestication of Pisum sativum： Is there a common genetic basis to the ‘domestication syndrome’ for legumes？ Annals of Botany， 2007， 100（5）： 1017-1025.
[62]	Jang S J， Sato M， Sato K， et al. A single-nucleotide polymorphism in an endo-1，4-β-glucanase gene controls seed coat permeability in soybean. PLoS One， 2015， 10（6）： e0128527.
[63]	Liu B， Fujita T， Yan Z H， et al. QTL mapping of domestication-related traits in soybean （Glycine max）. Annals of Botany， 2007， 100（5）： 1027-1038.
[64]	Chai M F， Zhou C E， Molina I， et al. A class Ⅱ KNOX gene， KNOX4， controls seed physical dormancy. Proceedings of the National Academy of Sciences， 2016， 113（25）： 6997-7002.
[65]	Chai M F， Castillo I Q， Sonntag A， et al. A seed coat-specific β-ketoacyl-CoA synthase， KCS12， is critical for preserving seed physical dormancy. Plant Physiology， 2021， 186（3）： 1606-1615.
[66]	Matilla A J. Seed dormancy： Molecular control of its induction and alleviation. Plants， 2020， 9（10）： 1402.
[67]	Koornneef M， Bentsink L， Hilhorst H. Seed dormancy and germination. Current Opinion in Plant Biology， 2002， 5（1）： 33-36.
[68]	Bassel G W. To grow or not to grow？ Trends in Plant Science， 2016， 21（6）： 498-505.
[69]	Shu K， Liu X， Xie Q， et al. Two faces of one seed： Hormonal regulation of dormancy and germination. Molecular Plant， 2016， 9（1）： 34-45.
[70]	Arc E， Sechet J， Corbineau F， et al. ABA crosstalk with ethylene and nitric oxide in seed dormancy and germination. Frontiers in Plant Science， 2013， 4： 63.
[71]	Hubbard K E， Nishimura N， Hitomi K， et al. Early abscisic acid signal transduction mechanisms： Newly discovered components and newly emerging questions. Genes & Development， 2010， 24（16）： 1695-1708.
[72]	Finkelstein R. Abscisic acid synthesis and response. The Arabidopsis Book， 2013， 11： e0166.
[73]	Cutler S R， Rodriguez P L， Finkelstein R R， et al. Abscisic acid： Emergence of a core signaling network. Annual Review of Plant Biology， 2010， 61（1）： 651-679.
[74]	Raghavendra A S， Gonugunta V K， Christmann A， et al. ABA perception and signalling. Trends in Plant Science， 2010， 15（7）： 395-401.
[75]	Ma Y， Szostkiewicz I， Korte A， et al. Regulators of PP2C phosphatase activity function as abscisic acid sensors. Science， 2009， 324（5930）： 1064-1068.
[76]	Park S Y， Fung P， Nishimura N， et al. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science， 2009， 324（5930）： 1068-1071.
[77]	Antoni R， Gonzalez-Guzman M， Rodriguez L， et al. Selective inhibition of clade a phosphatases type 2C by PYR/PYL/RCAR abscisic acid receptors. Plant Physiology， 2012， 158（2）： 970-980.
[78]	Umezawa T， Sugiyama N， Mizoguchi M， et al. Type 2C protein phosphatases directly regulate abscisic acid-activated protein kinases in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America， 2009， 106（41）： 17588-17593.
[79]	Gao Y L， Liu F，Xu F， et al. Saturated ceramide is required for seed germination in soybean. Seed Biology， 2025， 4（1）： e006.
[80]	Nakashima K， Fujita Y， Kanamori N， et al. Three Arabidopsis SnRK2 protein kinases， SRK2d/SnRK2.2， SRK2e/SnRK2.6/OST1 and SRK2i/SnRK2.3， involved in ABA signaling are essential for the control of seed development and dormancy. Plant and Cell Physiology， 2009， 50（7）： 1345-1363.
[81]	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. The Plant Cell， 2008， 20（10）： 2729-2745.
[82]	Fujii H， Zhu J K. Arabidopsis mutant deficient in 3 abscisic acid-activated protein kinases reveals critical roles in growth， reproduction， and stress. Proceedings of the National Academy of Sciences of the United States of America， 2009， 106（20）： 8380-8385.
[83]	van der Schaar W， Alonso-Blanco C， Léon-Kloosterziel K M， et al. QTL analysis of seed dormancy in Arabidopsis using recombinant inbred lines and MQM mapping. Heredity， 1997， 79（2）： 190-200.
[84]	Bentsink L， Jowett J， Hanhart C J， et al. Cloning of DOG1， a quantitative trait locus controlling seed dormancy in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America， 2006， 103（45）： 17042-17047.
[85]	Carrillo-Barral N， Rodríguez-Gacio M D C， Matilla A J. Delay of germination-1 （DOG1）： A key to understanding seed dormancy. Plants， 2020， 9（4）： 480.
[86]	Graeber K， Voegele A， Buttner-Mainik A， et al. Spatiotemporal seed development analysis provides insight into primary dormancy induction and evolution of the Lepidium delay of germination-1 genes. Plant Physiology， 2013， 161（4）： 1903-1917.
[87]	Tognacca R S， Botto J F. Post-transcriptional regulation of seed dormancy and germination： Current understanding and future directions. Plant Communications， 2021， 2（4）： 100169.
[88]	Chiang G C K， Bartsch M， Barua D， et al. DOG1 expression is predicted by the seed-maturation environment and contributes to geographical variation in germination in Arabidopsis thaliana. Molecular Ecology， 2011， 20（16）： 3336-3349.
[89]	Yatusevich R， Fedak H， Ciesielski A， et al. Antisense transcription represses Arabidopsis seed dormancy QTL DOG1 to regulate drought tolerance. EMBO Reports， 2017， 18（12）： 2186-2196.
[90]	Buijs G， Kodde J， Groot S P C， et al. Seed dormancy release accelerated by elevated partial pressure of oxygen is associated with DOG loci. Journal of Experimental Botany， 2018， 69（15）： 3601-3608.
[91]	Yang Y Z， Zheng C， Chandrasekaran U， et al. Identification and bioinformatic analysis of the GmDOG1-like family in soybean and investigation of their expression in response to gibberellic acid and abscisic acid. Plants， 2020， 9（8）： 937.
[92]	Dekkers B J W， He H Z， Hanson J， et al. The Arabidopsis DELAY of GERMINATION 1 gene affects ABSCISIC ACID INSENSITIVE 5 （ABI5） expression and genetically interacts with ABI3 during Arabidopsis seed development. The Plant Journal， 2016， 85（4）： 451-465.
[93]	Zinsmeister J， Lalanne D， Terrasson E， et al. ABI5 is a regulator of seed maturation and longevity in legumes. The Plant Cell， 2016， 28（11）： 2735-2754.
[94]	Carles C， Etheve N B， Aspart L， et al. Regulation of Arabidopsis thaliana Em genes： Role of ABI5. The Plant Journal， 2002， 30（3）： 373-383.
[95]	Molina L L， Mongrand S， McLachlin D T， et al. ABI5 acts downstream of ABI3 to execute an ABA-dependent growth arrest during germination. The Plant Journal， 2002， 32（3）： 317-328.
[96]	Zinsmeister J， Berriri S， Basso D P， et al. The seed-specific heat shock factor A9 regulates the depth of dormancy in Medicago truncatula seeds via ABA signalling. Plant， Cell & Environment， 2020， 43（10）： 2508-2522.
[97]	Kotak S， Vierling E， Umlein H B， et al. A novel transcriptional cascade regulating expression of heat stress proteins during seed development of Arabidopsis. The Plant Cell， 2007， 19（1）： 182-195.
[98]	Lalanne D， Malabarba J， Vu J L， et al. Medicago ABI3 splicing isoforms regulate the expression of different gene clusters to orchestrate seed maturation. Plants， 2021， 10（8）： 1710.
[99]	Li P， Ni H H， Ying S B， et al. Teaching an old dog a new trick： Multifaceted strategies to control primary seed germination by DELAY of GERMINATION 1 （DOG1）. Phyton， 2020， 89（1）： 1-12.
[100]	Iwasaki M， Penfield S， Lopez-Molina L. Parental and environmental control of seed dormancy in Arabidopsis thaliana. Annual Review of Plant Biology， 2022， 73（1）： 355-378.
[101]	Sajeev N， Koornneef M， Bentsink L. A commitment for life： Decades of unraveling the molecular mechanisms behind seed dormancy and germination. The Plant Cell， 2024， 36（5）： 1358-1376.
[102]	Penfield S， Josse E， Kannangara R， et al. Cold and light control seed germination through the bHLH transcription factor SPATULA. Current Biology， 2005， 15（22）： 1998-2006.
[103]	Rieu I， Eriksson S， Powers S J， et al. Genetic analysis reveals that c19-GA 2-oxidation is a major gibberellin inactivation pathway in Arabidopsis？ The Plant Cell， 2008， 20（9）： 2420-2436.
[104]	Rehmani M S， Aziz U， Xian B， et al. Seed dormancy and longevity： A mutual dependence or a trade-off？ Plant and Cell Physiology， 2022， 63（8）： 1029-1037.
[105]	Holdsworth M J， Bentsink L， Soppe W J J. Molecular networks regulating Arabidopsis seed maturation， after-ripening， dormancy and germination. New Phytologist， 2008， 179（1）： 33-54.
[106]	Tian R， Wang F F， Zheng Q L， et al. Direct and indirect targets of the Arabidopsis seed transcription factor ABSCISIC ACID INSENSITIVE3. The Plant Journal， 2020， 103（5）： 1679-1694.
[107]	Carbonero P， Iglesias-Fernández R， Vicente-Carbajosa J. The AFL subfamily of B3 transcription factors： Evolution and function in angiosperm seeds. Journal of Experimental Botany， 2017， 68（4）： 871-880.
[108]	Nishimura N， Tsuchiya W， 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）： 1-14
[109]	Bryant F M， Hughes D， Hassani-Pak K， et al. Basic LEUCINE ZIPPER TRANSCRIPTION FACTOR67 transactivates DELAY of GERMINATION1 to establish primary seed dormancy in Arabidopsis. The Plant Cell， 2019， 31（6）： 1276-1288.

类型Type	种子休眠原因Reasons for seed dormancy	文献References
PD	生理休眠一般由激素平衡来解释，其中脱落酸作为抑制剂，赤霉素作为促进剂。Physiological dormancy is typically elucidated through the lens of hormonal equilibrium， wherein abscisic acid （ABA） functions as an inhibitory regulator， while gibberellins （GA） serve as promotive agents.	［23］
PY	种子的不透水性导致种子不能吸收水分，因此不能发芽。The impermeability of the seed coat prevents water absorption， thereby inhibiting germination.	［33］
PY+PD	种子同时存在两种或多种休眠类型，既具有生理休眠的特点，内部含有抑制萌发的物质，又有物理休眠，种皮阻碍了种子与外界环境的物质交换。Seeds may exhibit multiple types of dormancy concurrently， characterized by both physiological dormancy， involving internal substances that inhibit germination， and physical dormancy， where the seed coat impedes material exchange with the external environment.	［37］

类型Type	种子休眠原因Reasons for seed dormancy	文献References
PD	生理休眠一般由激素平衡来解释，其中脱落酸作为抑制剂，赤霉素作为促进剂。Physiological dormancy is typically elucidated through the lens of hormonal equilibrium， wherein abscisic acid （ABA） functions as an inhibitory regulator， while gibberellins （GA） serve as promotive agents.	［23］
PY	种子的不透水性导致种子不能吸收水分，因此不能发芽。The impermeability of the seed coat prevents water absorption， thereby inhibiting germination.	［33］
PY+PD	种子同时存在两种或多种休眠类型，既具有生理休眠的特点，内部含有抑制萌发的物质，又有物理休眠，种皮阻碍了种子与外界环境的物质交换。Seeds may exhibit multiple types of dormancy concurrently， characterized by both physiological dormancy， involving internal substances that inhibit germination， and physical dormancy， where the seed coat impedes material exchange with the external environment.	［37］

解除技术 Release technology	种子休眠解除原理 Seed dormancy release mechanisms	解除类型 Categories of dormancy release
层积处理Stratification treatment	软化或分解种皮、促使种子内部发生复杂的生理生化变化。此外，完成胚的发育。Softening or decomposing the seed coat， promoting complex physiological and biochemical changes within the seed， and in addition， completing the development of the embryo.	PD， PY+PD
机械处理Mechanical treatment	物理外力破坏种子坚硬或致密的外皮以解除其对水分、气体渗透或胚生长的阻碍，从而打破休眠。Physical external forces destroy the hard or dense outer coat of seeds to remove their obstruction to water and gas penetration or embryo growth， thereby breaking dormancy.	PY
热水处理Hot water treatment	高温使种皮蜡质层破裂或产生微裂纹，增强透水性，热激改变种子内源激素平衡。High temperature causes the waxy layer of the seed coat to crack or form microcracks， enhances water permeability， and heat shock alters the balance of endogenous hormones in seeds.	PY， PD
激素处理Hormonal treatment	抑制脱落酸作用，促进赤霉素合成、刺激胚发育。Inhibit the action of abscisic acid（ABA）， promote the synthesis of gibberellin（GA）， and stimulate embryo development.	PD
硫酸处理Sulfuric acid treatment	降低种壳机械强度，破坏种皮的一些局部结构、改善种皮的渗透性、急剧降温导致种皮开裂，进而提升种皮通透性。Reduce the mechanical strength of the seed coat， destroy some local structures of the seed coat， improve the permeability of the seed coat， and cause the seed coat to crack due to rapid cooling， thereby enhancing the permeability of the seed coat.	PY
液氮处理Liquid nitrogen treatment	使种皮产生微裂纹，或改变细胞膜透性及酶活性。Cause the seed coat to form microcracks， or alter the permeability of cell membranes and enzyme activity.	PD， PY
超声波处理Ultrasonic wave treatment	利用超声波在液体中产生空化气泡，气泡破裂时释放冲击波，使种皮产生微孔或剥离外层屏障。Ultrasonic waves are used to generate cavitation bubbles in liquids； when the bubbles collapse， they release shock waves， causing the seed coat to form micropores or peel off the outer barrier.	PY

解除技术 Release technology	种子休眠解除原理 Seed dormancy release mechanisms	解除类型 Categories of dormancy release
层积处理Stratification treatment	软化或分解种皮、促使种子内部发生复杂的生理生化变化。此外，完成胚的发育。Softening or decomposing the seed coat， promoting complex physiological and biochemical changes within the seed， and in addition， completing the development of the embryo.	PD， PY+PD
机械处理Mechanical treatment	物理外力破坏种子坚硬或致密的外皮以解除其对水分、气体渗透或胚生长的阻碍，从而打破休眠。Physical external forces destroy the hard or dense outer coat of seeds to remove their obstruction to water and gas penetration or embryo growth， thereby breaking dormancy.	PY
热水处理Hot water treatment	高温使种皮蜡质层破裂或产生微裂纹，增强透水性，热激改变种子内源激素平衡。High temperature causes the waxy layer of the seed coat to crack or form microcracks， enhances water permeability， and heat shock alters the balance of endogenous hormones in seeds.	PY， PD
激素处理Hormonal treatment	抑制脱落酸作用，促进赤霉素合成、刺激胚发育。Inhibit the action of abscisic acid（ABA）， promote the synthesis of gibberellin（GA）， and stimulate embryo development.	PD
硫酸处理Sulfuric acid treatment	降低种壳机械强度，破坏种皮的一些局部结构、改善种皮的渗透性、急剧降温导致种皮开裂，进而提升种皮通透性。Reduce the mechanical strength of the seed coat， destroy some local structures of the seed coat， improve the permeability of the seed coat， and cause the seed coat to crack due to rapid cooling， thereby enhancing the permeability of the seed coat.	PY
液氮处理Liquid nitrogen treatment	使种皮产生微裂纹，或改变细胞膜透性及酶活性。Cause the seed coat to form microcracks， or alter the permeability of cell membranes and enzyme activity.	PD， PY
超声波处理Ultrasonic wave treatment	利用超声波在液体中产生空化气泡，气泡破裂时释放冲击波，使种皮产生微孔或剥离外层屏障。Ultrasonic waves are used to generate cavitation bubbles in liquids； when the bubbles collapse， they release shock waves， causing the seed coat to form micropores or peel off the outer barrier.	PY