藜麦穗发芽调控机制研究进展

doi:10.11924/j.issn.1000-6850.casb2025-0734

中国农学通报 ›› 2026, Vol. 42 ›› Issue (19): 60-66.doi: 10.11924/j.issn.1000-6850.casb2025-0734

藜麦穗发芽调控机制研究进展

陶红¹^,²(), 徐红卫¹, 宗营杰¹, 何婷¹, 刘成洪¹()

¹ 上海市农业科学院生物技术研究所/上海市农业遗传育种重点实验室，上海 201106
² 上海师范大学生命科学学院，上海 200234

收稿日期:2025-08-21 修回日期:2026-01-11 出版日期:2026-10-15 发布日期:2026-10-15
通讯作者:
刘成洪，男，1975年出生，湖北宜昌人，研究员，博士，研究方向：麦类作物遗传育种及功能开发研究。通信地址：201106 上海市闵行区北翟路2901号上海市农业科学院，Tel：021-62202965，E-mail：liuchenghong@saas.sh.cn。
作者简介:
陶红，女，2001年出生，安徽六安人，硕士研究生，研究方向：藜麦遗传育种。通信地址：201106 上海市闵行区北翟路2901号上海农业科学院生物所，Tel：021-62202965，E-mail：1000569353@smail.shnu.edu.cn。
基金资助:
上海市农业科技创新项目“藜麦小孢子单倍体育种关键技术研发与应用”[沪农科(T2024102)]; 2025年度关键技术研发计划“农业科技”项目“宜机化藜菜新品种培育”(25N22800300); 中国（上海）国际藜麦创新中心建设[农科国际2026(02)]

Research Progress on Regulatory Mechanisms of Pre-harvest Sprouting in Quinoa

TAO Hong¹^,²(), XU Hongwei¹, ZONG Yingjie¹, HE Ting¹, LIU Chenghong¹()

¹ Biotech Research Institute, Shanghai Academy of Agricultural Sciences/ Shanghai Key Laboratory of Agricultural Genetics and Breeding, Shanghai 201106
² College of Life Sciences, Shanghai Normal University, Shanghai 200234

Received:2025-08-21 Revised:2026-01-11 Published:2026-10-15 Online:2026-10-15

摘要/Abstract

摘要：

穗发芽是制约藜麦产量与品质提升的关键瓶颈。为深入理解藜麦穗发芽的调控机制并研发有效的防控策略，综述近5年藜麦穗发芽相关研究进展，重点分析种子生理特性、环境条件及植物激素等关键影响因素，总结其遗传调控基础与防控技术体系。研究表明，藜麦穗发芽受种子休眠特性、胚乳结构及环境温湿度相互作用的共同调控，多种植物激素通过信号网络调节种子休眠与萌发平衡，相关代谢的关键基因已初步解析，其中CqNCED等关键位点为培育抗穗发芽品种提供了重要的分子靶标。当前，多组学解析与生物技术应用已成为藜麦穗发芽研究的重要手段，但其关键调控机制仍有待深入解析，如环境—激素信号的耦合机制尚不清晰、特有休眠调控通路仍缺乏充分验证。未来需要加强基因编辑、单倍体育种等先进育种技术应用，推动藜麦抗穗发芽等优良新种质创制，为藜麦产业高质量发展提供技术支撑。

关键词: 藜麦, 穗发芽, 植物激素, 分子机制, 防控策略

Abstract:

Pre-harvest sprouting (PHS) is a major constraint limiting the improvement of quinoa yield and quality. To enhance the understanding of the regulatory mechanisms underlying PHS in quinoa and advance research on management strategies, this study reviews the progress on PHS in quinoa over the past five years. Key influencing factors such as seed physiological characteristics, environmental conditions, and phytohormones are discussed, and the genetic regulatory basis and technical approaches for PHS control are outlined. Studies indicate that quinoa PHS is co-regulated by seed dormancy characteristics, endosperm structure, and interactions between ambient temperature and humidity. Various phytohormones coordinate the balance between dormancy and germination through signaling networks, and key genes involved in related metabolic pathways have been preliminarily identified. Key loci such as CqNCED provide important molecular targets for breeding pre-harvest sprouting-resistant varieties. Although multi-omics approaches and biotechnological applications have become important tools in studies on PHS in quinoa, the key regulatory mechanisms still require further elucidation. For instance, the coupling mechanisms between environmental and hormone signaling remain unclear, and the dormancy regulatory pathways specific to quinoa are still not fully validated. Future efforts should strengthen the application of advanced breeding technologies such as gene editing and haploid breeding to promote the development of new quinoa germplasm with enhanced resistance to pre-harvest sprouting, thereby providing technical support for the sustainable development of the quinoa industry.

Key words: quinoa, pre-harvest sprouting, phytohormones, molecular mechanism, control strategy

中图分类号:

S519其他

陶红, 徐红卫, 宗营杰, 何婷, 刘成洪. 藜麦穗发芽调控机制研究进展[J]. 中国农学通报, 2026, 42(19): 60-66.

TAO Hong, XU Hongwei, ZONG Yingjie, HE Ting, LIU Chenghong. Research Progress on Regulatory Mechanisms of Pre-harvest Sprouting in Quinoa[J]. Chinese Agricultural Science Bulletin, 2026, 42(19): 60-66.

图/表 1

参考文献 58

[1]	PATHAN S, SIDDIQUI R A. Nutritional composition and bioactive components in quinoa (Chenopodium quinoa Willd.) greens: A review[J]. Nutrients, 2022, 14(3):558. doi: 10.3390/nu14030558 URL
[2]	姜庆国, 温日宇, 郭耀东, 等. 藜麦的种植栽培技术与病虫害防治[J]. 农民致富之友, 2017(22):154.
[3]	TABATABAEI I, ALSEEKH S, SHAHID M, et al. The diversity of quinoa morphological traits and seed metabolic composition[J]. Scientific data, 2022, 9(1):323. doi: 10.1038/s41597-022-01399-y pmid: 35725573
[4]	SHA H, YU Y, HAN Y, et al. Combination of maleic hydrazide and coumarin inhibits rice seed germination involving reactive oxygen species accumulation, ABA metabolism and starch degradation[J]. Plant physiology and biochemistry, 2024, 217:109202. doi: 10.1016/j.plaphy.2024.109202 URL
[5]	WU Q, BAI X, WU X, et al. Transcriptome profiling identifies transcription factors and key homologs involved in seed dormancy and germination regulation of Chenopodium quinoa[J]. Plant physiology and biochemistry, 2020, 151:443-456. doi: S0981-9428(20)30163-7 pmid: 32289638
[6]	杨招娣, 郭凤根, 王仕玉, 等. 植物生长抑制剂对藜麦农艺性状和穗发芽抗性的影响[J]. 中国农业科技导报, 2024, 26(5):44-51.
[7]	林鑫焱, 张传忠, 戴兵, 等. 水稻穗发芽遗传与分子机制的研究进展[J]. 生物技术通报, 2024, 40(1):24-31. doi: 10.13560/j.cnki.biotech.bull.1985.2023-0786
[8]	MCGINTY E M, MURPHY K M, HAUVERMA L, et al. Seed dormancy and preharvest sprouting in quinoa (Chenopodium quinoa Willd.)[J]. Plants (Basel), 2021, 10(3):458.
[9]	AGÜERO-MARTÍNEZ P F, CARDOZO L, GÓMEZ C A, et al. Variation in thickness of embryo covering structures and their role in the regulation of seed physiological dormancy of Chenopodium hircinum (Amaranthaceae)[J]. Plants (Basel), 2024, 13(19):2832.
[10]	MCGINTY E M, CRAINE E B, MILLER N D, et al. Evaluating relationships between seed morphological traits and seed dormancy in Chenopodium quinoa Willd[J]. Frontiers in plant science, 2023, 14:1161165. doi: 10.3389/fpls.2023.1161165 URL
[11]	王慧丽, 钟苑宁, 杨晗, 等. 水分和温度对藜麦种子萌发出苗及籽粒产量的影响[J]. 中国生态农业学报(中英文), 2025, 33(1):95-103.
[12]	毛琪, 晏兴珠, 王仕玉, 等. 30份藜麦资源的穗发芽抗性评价[J]. 种子, 2021, 40(10):62-66,73.
[13]	朱丽丽, 张发玉, 安宁, 等. 116份藜麦种质资源萌发期抗旱性综合评价[J]. 干旱地区农业研究, 2024, 42(1):23-31.
[14]	高艳梅, 冯鹏睿, 陈薇薇, 等. 抗旱性藜麦幼苗对干旱胁迫的生理响应[J]. 作物杂志, 2026(1):182-188.
[15]	卢秋巍, 王媛媛, 辛宇轩, 等. 冷、旱胁迫及ABA、氟啶酮处理下藜麦qPCR内参基因的筛选[J]. 农业生物技术学报, 2024, 32(12):2891-2903.
[16]	王悦娟, 赵延蓉, 魏玉清. 不同温度条件下盐、碱和干旱胁迫对藜麦种子萌发过程的影响[J]. 分子植物育种, 2025, 23(2):543-553.
[17]	曲波, 张谨华, 王鑫, 等. 温度和光照对藜麦幼苗生长发育的影响[J]. 农业工程, 2018, 8(7):128-131.
[18]	廖映秀, 戴红燕, 杨蛟, 等. 温度对陇藜1号种子萌发和幼苗生长的影响[J]. 安徽农业科学, 2020, 48(19):30-33.
[19]	宿婧, 梁彬, 郝晓华, 等. 外源褪黑素对干旱胁迫下黑藜麦种子萌发及生理特性的影响[J]. 江苏农业科学, 2025, 53(6):125-129.
[20]	李凤, 郭元元, 刘高洋, 等. 外源ALA对盐胁迫和干旱胁迫下藜麦种子萌发的缓解效应[J]. 种子, 2025, 44(1):188-196,203.
[21]	赵颖, 魏小红, 赫亚龙, 等. 混合盐碱胁迫对藜麦种子萌发和幼苗抗氧化特性的影响[J]. 草业学报, 2019, 28(2):156-167. doi: 10.11686/cyxb2018181
[22]	王洋, 杜会石, 鲍庆晗. 藜麦种质资源萌发期抗旱性综合评价及抗旱指标筛选[J]. 江苏农业科学, 2023, 51(19):62-68.
[23]	杜晨曦, 王金丽, 周华坤, 等. 赤霉素对植物种子萌发及幼苗生长影响的研究进展[J]. 湖北农业科学, 2019, 58(22):9-14.
[24]	蒋梦珂, 谷林珠, 马艳雨, 等. 赤霉素浸种对藜麦种子萌发及生理特性的影响[J]. 天津农业科学, 2024, 30(11):1-7.
[25]	GAO Y, ZHENG C, GE W, et al. Integrating transcriptome, physiological, and biochemical studies revealing the role of endogenous ABA and GA₃ in the germination of quinoa seed[J/OL]. Research square, 2024-10-29[2026-03-05]. https://doi.org/10.21203/rs.3.rs-5283572/v1.
[26]	NAZIH A, BAGHOUR M, MAATOUGUI A, et al. Effect of gibberellic acid and mechanical scarification on the germination and seedling stages of Chenopodium quinoa Willd. under salt stress[J]. Plants (Basel), 2024, 13(10):1330.
[27]	王德领, 庄东英, 蒋润枝, 等. 外源ABA对藜麦种子萌发的影响[J]. 中国种业, 2022(2):93-96.
[28]	操春燕, 陈熙, 吴燕. 脱落酸调控种子萌发和休眠的研究进展[J]. 生命科学, 2017, 29(6):582-588.
[29]	ZENG F, ZHENG C, GE W, et al. Regulatory function of the endogenous hormone in the germination process of quinoa seeds[J]. Frontiers in plant science, 2024, 14:1322986. doi: 10.3389/fpls.2023.1322986 URL
[30]	陈凡, 丰扬, 王仕玉, 等. 外源ABA对藜麦种子萌发及ABA相关基因表达的影响[J]. 西北植物学报, 2023, 43(2):295-304.
[31]	骞帆, 曾芳, 高娅, 等. 不同外源激素对藜麦发芽特性的影响[J]. 分子植物育种, 2024.
[32]	景茂雅, 张子玉, 张萌, 等. 水杨酸浸种对盐胁迫藜麦种子萌发及幼苗生长的影响[J]. 作物杂志, 2025(1):194-201.
[33]	HAO Y, HONG Y, GUO H, et al. Transcriptomic and metabolomic landscape of quinoa during seed germination[J]. Biomed central plant biology, 2022, 22(1):237.
[34]	YANG F, WANG S Y, GUO F G, et al. Identification and characterization of the Chenopodium quinoa gibberellin oxidase gene family and its role in seed germination[J]. South African journal of botany, 2025, 176:219-230. doi: 10.1016/j.sajb.2024.11.027 URL
[35]	OKAMOTO M, KUWAHARA A, SEO M, et al. CYP707A1 and CYP707A2, which encode abscisic acid 8’-hydroxylases, are indispensable for proper control of seed dormancy and germination in Arabidopsis[J]. Plant physiology, 2006, 141(1):97-107. doi: 10.1104/pp.106.079475 URL
[36]	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[J]. 中国科学:生命科学, 2020, 50(6):599-615.
[37]	FENG Y, GUO F, WANG S, et al. Identification and functional analysis of the CqGID1 homologs in quinoa[J]. Plant cell reports, 2025, 44:192. doi: 10.1007/s00299-025-03579-7 pmid: 40762808
[38]	HUANG L, ZHANG L, ZHANG P, et al. Comparative transcriptomes and WGCNA reveal hub genes for spike germination in different quinoa lines[J]. Biomed central genomics, 2024, 25(1):1231.
[39]	MANZOOR N, YUAN J, DONGCHENG W, et al. Integrated transcriptomic and proteomic analyses revealed molecular mechanisms underlying nutritional changes during seed development of Chenopodium quinoa[J]. Genomics, 2025, 117(3):111045. doi: 10.1016/j.ygeno.2025.111045 URL
[40]	王春妹, 王梅, 王红霞, 等. 干旱胁迫下藜麦种子糖代谢转录组学研究[J]. 植物遗传资源学报, 2024, 25(8):1370-1384.
[41]	WANG C, LU C, WANG J, et al. Molecular mechanisms regulating glucose metabolism in quinoa (Chenopodium quinoa Willd.) seeds under drought stress[J]. Biomed central plant biology, 2024, 24(1):796.
[42]	YAN H, ZHANG Z, LV Y, et al. Integrated multispectral imaging, germination phenotype, and transcriptomic analysis provide insights into seed vigor responsive mechanisms in quinoa under artificial accelerated aging[J]. Frontiers in plant science, 2024, 15:1435154. doi: 10.3389/fpls.2024.1435154 URL
[43]	张娜, 罗于洋, 马迎梅, 等. 盐胁迫对藜麦种子萌发的影响研究[J]. 广西植物, 2022, 42(6):1029-1039.
[44]	ZHU X, WANG B, LIU W, et al. Genome-wide analysis of AP2/ERF gene and functional analysis of CqERF24 gene in drought stress in quinoa[J]. International journal of biological macromolecules, 2023(8):127582.
[45]	刘文瑜, 李健荣, 杨发荣, 等. 盐胁迫对40份藜麦种质种子萌发的影响及其耐盐性评价[J]. 分子植物育种, 2021(3):983-995.
[46]	毋悦悦, 庞春花, 张永清, 等. 壳聚糖引发对盐胁迫下藜麦种子萌发和幼苗生长的影响[J]. 江苏农业科学, 2024, 52(4):114-121.
[47]	杨利艳, 杨小兰, 朱满喜, 等. 干旱胁迫对藜麦种子萌发及幼苗生理特性的影响[J]. 种子, 2020(9):36-40.
[48]	MA L, DING S, FU X, et al. Enzymatic and transcriptomic analysis reveals the essential role of carbohydrate metabolism in freesia (Freesia hybrida) corm formation[J]. PeerJ, 2021, 9:e11078. doi: 10.7717/peerj.11078 URL
[49]	李晟涵, 崔立刚, 徐超, 等. 北京市通州区藜麦种植及田间管理技术[J]. 河南农业, 2025(6):28-30.
[50]	高东尧, 许纪发, 李臻, 等. 小麦穗发芽研究进展[J]. 湖南农业科学, 2024(11):107-112.
[51]	FU R, LIANG X, LI J, et al. Comparative transcriptomic analyses reveal key pathways in response to cold stress at the germination stage of quinoa (Chenopodium quinoa Willd.) Seeds[J]. Plants (Basel), 2025, 14(8):1212.
[52]	CECCATO D, BERTERO D, BATLLA D, et al. Structural aspects of dormancy in quinoa (Chenopodium quinoa): Importance and possible action mechanisms of the seed coat[J]. Seed science research, 2015, 25:267-275. doi: 10.1017/S096025851500015X URL
[53]	BARAKAT H, AL-QABBA M M, ALGONAIMAN R, et al. Impact of sprouting process on the protein quality of yellow and red quinoa (Chenopodium quinoa)[J]. Molecules, 2024, 29(2):404. doi: 10.3390/molecules29020404 URL
[54]	PAN W, LIANG J, SUI J, et al. ABA and bud dormancy in perennials: Current knowledge and future perspective[J]. Genes (Basel), 2021, 12(10):1635. doi: 10.3390/genes12101635 URL
[55]	OCAÑA-GALLEGOS C, LIANG M, MCGINTY E, et al. Preharvest sprouting in quinoa: A new screening method adapted to panicles and GWAS components[J]. Plants (Basel), 2024, 13(10):1297.
[56]	BU G, MA X, JIN H, et al. CRISPR-Cas and RNA sequencing reveal nutrient enhancement pathways in quinoa for plant-based athlete recovery diets and future food security[J]. International journal of biological macromolecules, 2025(4):144785.
[57]	QU Y, FERNIE A R, LIU J, et al. Doubled haploid technology and synthetic apomixis: Recent advances and applications in future crop breeding[J]. Molecular plant, 2024, 17(7):1005-1018. doi: 10.1016/j.molp.2024.06.005 URL
[58]	李玲红, 苟彤, 任爱霞, 等. 藜麦基因组学与重要农艺性状位点研究进展[J]. 遗传, 2022, 44(11):1009-1027.

基因名称	注释	基因序列号	参考文献
CqCYP707A1	Abscisic Acid 8’-hydroxylase	AUR62030408	[33]
CqNCED	9-cis-Epoxycarotenoid Dioxygenase	XM_021893846.1	[33]
CqGID1	Gibberellin Insensitive Dwarf1	LOC110699090	[33]
CqABA1	Zeaxanthin epoxidase	AUR62001926	[33]
CqGA20ox	Gibberellin 20-oxidase	LOC110726394	[35]
CqGA3ox	Gibberellin 3β-dioxygenase	LOC110701233	[35]
CqGA2ox_2	Gibberellin 2β-Dioxygenase	AUR62011753	[35]
CqGA2ox_1	Gibberellin 2β-Dioxygenase	AUR62024597	[35]
CqGID1b1	Gibberellin Insensitive Dwarf1b, isoform 1	AUR62000900	[36]
CqGID1b2	Gibberellin Insensitive Dwarf1b, isoform 2	AUR62034591	[36]
CqAMY1	α-amylase 1	AUR62012979	[39]
CqBAM3	β-amylase 3	AUR62007199	[39]
CqINV	Invertase	LOC110719866	[40]
CqFrK	Fructokinase	LOC110734349
CqBGAL	β-galactosidase	AUR62011825	[41]

基因名称	注释	基因序列号	参考文献
CqCYP707A1	Abscisic Acid 8’-hydroxylase	AUR62030408	[33]
CqNCED	9-cis-Epoxycarotenoid Dioxygenase	XM_021893846.1	[33]
CqGID1	Gibberellin Insensitive Dwarf1	LOC110699090	[33]
CqABA1	Zeaxanthin epoxidase	AUR62001926	[33]
CqGA20ox	Gibberellin 20-oxidase	LOC110726394	[35]
CqGA3ox	Gibberellin 3β-dioxygenase	LOC110701233	[35]
CqGA2ox_2	Gibberellin 2β-Dioxygenase	AUR62011753	[35]
CqGA2ox_1	Gibberellin 2β-Dioxygenase	AUR62024597	[35]
CqGID1b1	Gibberellin Insensitive Dwarf1b, isoform 1	AUR62000900	[36]
CqGID1b2	Gibberellin Insensitive Dwarf1b, isoform 2	AUR62034591	[36]
CqAMY1	α-amylase 1	AUR62012979	[39]
CqBAM3	β-amylase 3	AUR62007199	[39]
CqINV	Invertase	LOC110719866	[40]
CqFrK	Fructokinase	LOC110734349
CqBGAL	β-galactosidase	AUR62011825	[41]

藜麦穗发芽调控机制研究进展

Research Progress on Regulatory Mechanisms of Pre-harvest Sprouting in Quinoa

RichHTML

PDF (PC)

可视化

摘要/Abstract

引用本文

使用本文

图/表 1

参考文献 58

相关文章 15

编辑推荐

Metrics

本文评价

关于本刊

作者中心

审者中心

在线期刊

联系我们

[1]	于洋洋, 张长远, 张惠尧, 吴廷全, 谢琦, 郭金菊. 葫芦科作物单性结实研究进展[J]. 中国农学通报, 2026, 42(5): 55-61.
[2]	王生萍, 卓玛曲措, 降志兵, 谈建鑫, 同坚. 西藏藜麦种子可培养内生细菌多样性[J]. 中国农学通报, 2026, 42(18): 25-32.
[3]	李娟, 李健, 马鹏飞, 杨庆利. 藜麦营养特征、烹饪特性及发展态势研究[J]. 中国农学通报, 2026, 42(10): 211-218.
[4]	汪锦, 万斌, 涂勋良, 蒋彧, 邓家洪. 兰花花色素沉着调控机理的研究进展[J]. 中国农学通报, 2026, 42(1): 84-91.
[5]	赵桂龙, 唐佳琦, 黄烁淇, 李荣田, 卜庆云, 鲁振强. 植物冷胁迫下ICE-CBF-COR耐冷调控分子机理的研究进展[J]. 中国农学通报, 2025, 41(9): 132-139.
[6]	苏芸芸, 徐梦馨, 欧晓彬, 龚磊, 张魁, 郭倩. 外源硒响应植物高温胁迫机制研究进展[J]. 中国农学通报, 2025, 41(33): 113-120.
[7]	李金涛, 佘奎军, 吴瑞, 杨国虎. 玉米矮秆基因研究进展[J]. 中国农学通报, 2025, 41(32): 9-16.
[8]	张鑫, 郭志鸿, 柴娟, 马荣, 周锋, 王琼, 姚卓男. 兰州百合组培增殖系数与鳞茎膨大的关键影响因子研究[J]. 中国农学通报, 2025, 41(31): 41-46.
[9]	廖慧, 张金凤, 张华, 张西西, 宋利娜, 张华丽, 舒健骅. 月季花器官变异的研究进展[J]. 中国农学通报, 2025, 41(19): 49-55.
[10]	关玉熙, 温超. 外源独脚金内酯对波斯菊种子萌发的影响[J]. 中国农学通报, 2025, 41(16): 40-44.
[11]	詹嘉涛, 缪乐怡, 纪子贤, 王一婷, 卢月, 谭恩, 马帅鹏, 王丽敏. 水稻OsSIG5基因对植物激素和非生物胁迫的响应分析[J]. 中国农学通报, 2025, 41(14): 32-40.
[12]	王政, 尹航, 张豪杰, 曾洋, 陈紫岩, 庞春霞, 朱娅玫, 王宏宝, 吴传万. 有机硒处理对藜麦菜硒强化及其产量品质的影响[J]. 中国农学通报, 2025, 41(13): 56-64.
[13]	张述伟, 黄琳丽, 宗营杰, 郭慧敏, 郭桂梅, 杨邦伟, 陈志伟, 徐红卫, 刘成洪. 藜麦苗总黄酮提取优化及其对黄嘌呤氧化酶的抑制作用和抗氧化活性评价[J]. 中国农学通报, 2025, 41(12): 142-148.
[14]	刘佳, 黄大荣, 尧美英, 刘硕, 张玉萍, 张国薇. 2个杏栽培品种需冷量差异的分子机制探索[J]. 中国农学通报, 2025, 41(1): 33-41.
[15]	江珊, 吴龙英, 赵宝生, 黄佳惠, 蒋宇喆, 焦元, 黄进. 植物耐受高温胁迫的分子机制研究进展[J]. 中国农学通报, 2024, 40(9): 132-138.