Article(id=1309574784824999997, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, articleNumber=null, orderNo=null, doi=10.11733/j.issn.1007-0435.2026.07.006, pmid=null, cstr=null, oa=null, hot=0, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1755705600000, receivedDateStr=2025-08-21, revisedDate=1760371200000, revisedDateStr=2025-10-14, acceptedDate=null, acceptedDateStr=null, onlineDate=1790156559425, onlineDateStr=2026-09-23, pubDate=null, pubDateStr=null, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1790156559425, onlineIssueDateStr=2026-09-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1790156559425, creator=13701087609, updateTime=1790156559425, updator=13701087609, issue=Issue{id=1309574765791244449, tenantId=1146029695717560320, journalId=1308397196074385509, year='2026', volume='34', issue='7', pageStart='2341', pageEnd='2752', issueExtLink='null', onlineDate='null', pubDate='1784044800000', pubDateStr='2026-07-15', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1790156554887, creator='13701087609', updateTime=1790156849638, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1309576002150424823, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1309576002154619128, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2403, endPage=2414, ext={EN=ArticleExt(id=1309574787324805183, articleId=1309574784824999997, tenantId=1146029695717560320, journalId=1308397196074385509, language=EN, title=Physiological Changes and Transcriptomic Analysis of Caucasian Clover under Low Temperature Stress, columnId=null, journalTitle=Acta Agrestia Sinica, columnName=null, runingTitle=null, highlight=null, articleAbstract=This study investigated the physiological and molecular mechanisms underlying the cold tolerance of Caucasian clover (Trifolium ambiguum M. Bieb.), a perennial forage known for its strong hardiness. Seedlings grown at 20℃ served as the control (CK), and low-temperature stress (4℃) was applied for 2, 6, and 12 h. Physiological indicators were measured at each time point, and transcriptome sequencing was conducted on the control and 12 h-stressed plants. The results indicated that under 4℃, the plants enhanced their cold tolerance by significantly increasing the activities of antioxidant enzymes (peroxidase, catalase, and superoxide dismutase) and the levels of osmoregulatory substances (soluble protein, soluble sugar, and free proline). This coordinated response effectively scavenged reactive oxygen species and mitigated membrane lipid peroxidation, as evidenced by reduced malondialdehyde accumulation. Transcriptome analysis identified 15 428 differentially expressed genes (DEGs) after 12 h of cold stress, with 8909 up-regulated and 6519 down-regulated. Key pathways were significantly activated, including the flavonoid biosynthesis pathway, where the up-regulation of pivotal upstream genes (CHS and CHI) facilitated the synthesis of antioxidant compounds such as luteolin, apigenin, and isoflavones. Concurrently, the MAPK signaling pathway was induced, highlighted by the increased expression of genes like MEKK1, suggesting that cold signals are transduced via a phosphorylation cascade. These findings provide crucial insights into the molecular basis of cold adaptation in Caucasian clover., authors=WU Qian1, ZHAO Ya-ru2, CAO Ke-fan1, MA Yi-ming1, SUO Yuan1, LI Shu-ning1, QI Hai-bo1, WANG Ming-jiu1, authorsList=WU Qian, ZHAO Ya-ru, CAO Ke-fan, MA Yi-ming, SUO Yuan, LI Shu-ning, QI Hai-bo, WANG Ming-jiu, authorCompany=1 College of Grassland Science, Inner Mongolia Agricultural University/Key Laboratory of Grassland Resource of Ministry of Education, Hohhot, Inner Mongolia 010011, China; 2 Inner Mongolia Zhengshi Ecological Agriculture (Group) Co., Ltd., Hohhot, Inner Mongolia 010010, China, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1309574786926346302, articleId=1309574784824999997, tenantId=1146029695717560320, journalId=1308397196074385509, language=CN, title=基于生理与转录组学的高加索三叶草耐寒性分析, columnId=1309565024058835950, journalTitle=草地学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=本研究以耐寒性强的多年生牧草高加索三叶草(Trifolium ambiguum M. Bieb.)为材料,以20℃正常生长的幼苗为对照(CK),设置4℃低温胁迫2 h,6 h 和 12 h共3个处理,测定各时期高加索三叶草的生理指标,并对CK与胁迫12 h的植株进行转录组测序,解析其幼苗期在低温胁迫下的生理反应与适应机制。结果表明,4℃处理下,植株通过提高抗氧化酶(过氧化物酶、过氧化氢酶、超氧化物歧化酶)活性及渗透调节物质(可溶性蛋白、可溶性糖、游离脯氨酸)含量,有效清除活性氧,减轻了膜脂过氧化损伤,从而增强其抗寒能力。转录组分析显示,4℃处理12 h后共鉴定到15 428个差异表达基因,其中8909个基因上调,6519个基因下调。其中类黄酮生物合成通路与MAPK信号通路被显著激活:类黄酮通路上游关键基因查尔酮合成酶和查尔酮异构酶显著上调,促进了抗氧化物质(木犀草素、芹菜素、异黄酮)的合成;MAPK通路中MEKK1等基因的表达升高,表明低温信号通过磷酸化级联反应进行转导。其研究结果为解析高加索三叶草耐寒性提供了理论依据。, authors=吴倩1, 赵雅茹2, 曹克璠1, 马一鸣1, 索媛1, 李舒宁1, 齐海波1, 王明玖1, authorsList=吴倩, 赵雅茹, 曹克璠, 马一鸣, 索媛, 李舒宁, 齐海波, 王明玖, authorCompany=1 内蒙古农业大学草业学院/草地资源教育部重点实验室, 内蒙古 呼和浩特 010011; 2 内蒙古正时生态农业(集团)有限公司, 内蒙古 呼和浩特 010010, correspAuthors=null, authorNote=吴倩(1999-),女,汉族,内蒙古丰镇人,硕士研究生,主要从事草地资源生态与管理研究,E-mail:15548302887@163.com, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=6zPbTiMcTCnHMvAkAHQmgw==, pdfFileSize=3162709, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=国家自然科学基金项目“蒙农三叶草1号结瘤固氮机理及根瘤菌多样性研究” (31960353); “蒙农三叶草1号花期延长及其调控机理研究” (32160334))}, authors=null, keywords=[Keyword(id=1309574849601822892, tenantId=1146029695717560320, journalId=1308397196074385509, articleId=1309574784824999997, language=CN, orderNo=1, keyword=高加索三叶草), Keyword(id=1309574849660543149, tenantId=1146029695717560320, journalId=1308397196074385509, articleId=1309574784824999997, language=CN, orderNo=2, keyword=低温胁迫), Keyword(id=1309574849937367214, tenantId=1146029695717560320, journalId=1308397196074385509, articleId=1309574784824999997, language=CN, orderNo=3, keyword=生理指标), Keyword(id=1309574849996087471, tenantId=1146029695717560320, journalId=1308397196074385509, articleId=1309574784824999997, language=CN, orderNo=4, keyword=差异基因), Keyword(id=1309574850121916592, tenantId=1146029695717560320, 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Effects of different temperatures on photosynthetic characteristics and expression of photosynthesis-related genes in amaranth[J]. Journal of Plant Resources and Environment, 2022, 31(5):50-57 郑友峰, 陈家兰, 肖昉, 等 不同温度对苋菜光合特性及光合作用相关基因表达的影响[J]. 植物资源与环境学报, 2022, 31(5):50-57 DING Y L, SHI Y T, YANG S H. Advances and challenges in uncovering cold tolerance regulatory mechanisms in plants[J]. New phytologist, 2019, 222(4):1690-1704 XIONG H Y, DUAN R J, WANG R S. Advances in physiological basis and molecular mechanisms of cold acclimation affecting photosynthetic characteristics and plant type characteristics in overwintering crops[J]. Crops, 2017, (4):21-26 熊辉岩, 段瑞君, 王瑞生. 冷驯化影响越冬作物光合特性和株型特征的生理基础与分子机制的研究进展[J]. 作物杂志, 2017, (4):21-26 XIANG H T, ZHENG D F, HE N, et al. Research progress on physiological responses of plants to low temperature stress and mitigation effects of exogenous abscisic acid[J]. Acta Prataculturae Sinica, 2021, 30(1):208-219 项洪涛, 郑殿峰, 何宁, 等. 植物对低温胁迫的生理响应及外源脱落酸缓解胁迫效应的研究进展[J]. 草业学报, 2021, 30(1):208-219 HUANG L F, LONG Y Z, LI J Q, et al. Effects of low temperature stress on physiological characteristics of Coffea arabica L. seedlings[J]. Journal of Agricultural Science and Technology, 2023, 25(2):60-67 黄丽芳, 龙宇宙, 李金芹, 等. 低温胁迫对小粒种咖啡幼苗生理特性的影响[J]. 中国农业科技导报, 2023, 25(2):60-67 WEI X X, LI Y M, LI Y B, et al. Correlation between cold resistance and phloem sugar content and endogenous hormone accumulation in grape[J]. Acta Agriculturae Boreali-occidentalis Sinica, 2021, 30(3):386-394 韦霞霞, 李玉梅, 李彦彪, 等. 葡萄抗寒性与韧皮部糖分及内源激素积累的相关性[J]. 西北农业学报, 2021, 30(3):386-394 YANG H Y, LIU C Z. Effects of low temperature stress on cold resistance physiological indexes of four Euonymus plants[J]. Grassland and Turf, 2020, 40(2):92-98 杨航宇, 刘长仲. 低温胁迫对4种卫矛属植物抗寒生理指标的影响[J]. 草原与草坪, 2020, 40(2):92-98 TAN H Q, HUANG H T, TIE M M, et al. Transcriptome profiling of two asparagus bean (Vigna unguiculata subsp. sesquipedalis) cultivars differing in chilling tolerance under cold stress[J]. PLoS One, 2017, 11(3):e0151105 SHEAFFER C C, MARTEN G C. Kura clover forage yield, forage quality, and stand dynamics[J]. Canadian Journal of Plant Science, 1991, 71(4):1169-1172 TOWNSEND C E. Phenotypic diversity for agronmic charaaters and frequency of self-compatible plants in Trifolum ambiguum[J]. Canadian Journal of Plant Science, 1970, 50(3):331-338 SPEER G S, ALLINSON D W. Kura clover (Trifolium ambiguum): Legume for forage and soil conservation[J]. Economic Botany, 1985, 39(2):165-176 YIN X J. Study on adaptability and utilization value of Trifolium ambiguum M. Bieb. in Inner Mongolia[D]. Hohhot: Inner Mongolia Agricultural University, 2004:5-9 殷秀杰. 高加索三叶草在内蒙古地区适应性和利用价值研究[D]. 呼和浩特:内蒙古农业大学, 2004:5-9 MARSHALL A H, RASCLE C, ABBERTON M T, et al. Introgression as a route to improved drought tolerance in white clover (Trifolium repens L.)[J]. Journal of Agronomy and Crop Science, 2001, 187(1):11-18 NGUYEN T D, ANDREWS M, LEY T W S,, et al. Caucasian clover (Trifolium ambiguum) specific rhizobia persist in low and high fertility soils in the South Island of New Zealand[J]. New Zealand Journal of Agricultural Research, 2020, 63(3):332-340 PENG X, SHI F L, FU N N. Relationship between pigment, polyphenol contents and leaf color parameters in Trifolium lupinaster leaves[J]. Acta Agrestia Sinica, 2023, 31(7):2005-2011 彭新, 石凤翎, 付娜娜. 野火球叶片色素和多酚类物质含量与叶色参数的关系[J]. 草地学报, 2023, 31(7):2005-2011 ZHUANG D F, WANG C, BAO J H, et al. Antioxidant enzyme activities and related gene expressions of Rosa xanthina F. normalis under low temperature stress[J]. Journal of Northeast Forestry University, 2023, 51(4):89-92 庄得凤, 王成, 包金花, 等. 低温胁迫下单瓣黄刺玫抗氧化酶活性及相关基因表达[J]. 东北林业大学学报, 2023, 51(4):89-92 WEN R Y, LIU J X, LI S, et al. Effects of low temperature stress on physiological and biochemical characteristics of different quinoa seedlings[J]. Seed, 2019, 38(5):53-56 温日宇, 刘建霞, 李顺, 等. 低温胁迫对不同藜麦幼苗生理生化特性的影响[J].种子, 2019, 38(5):53-56 SUN Y L, LI J D, SUN B, et al. Response of main cold resistance physiological indexes of different alfalfa varieties to low temperature[J]. Journal of Shenyang Agricultural University, 2017, 48(5):591-596 孙予璐, 李建东, 孙备, 等. 不同品种紫花苜蓿主要抗寒生理指标对低温的响应[J]. 沈阳农业大学学报, 2017, 48(5):591-596 GAO T T, MAO P C, TIAN X X, et al. Evaluation of cold tolerance of 10 white clover varieties at seedling stage[J]. Pratacultural Science, 2014, 31(9):1724-1731 高婷婷, 毛培春, 田小霞, 等. 10个白三叶品种苗期耐寒性评价[J]. 草业科学, 2014, 31(9):1724-1731 WU Y X, CAI C Y, TANG S B, et al. Research progress on growth, development and molecular mechanisms of plants in response to low temperature[J]. Jiangsu Agricultural Sciences, 2023, 51(19):1-9 吴宇欣, 蔡昌杨, 唐诗蓓, 等. 植物响应低温的生长发育及分子机制研究进展[J]. 江苏农业科学, 2023, 51(19):1-9 SU L W, LI S, MA S Y, et al. Establishment of a comprehensive evaluation method for grape cold resistance[J]. Acta Prataculturae Sinica, 2015, 24(3):70-79 苏李维, 李胜, 马绍英, 等. 葡萄抗寒性综合评价方法的建立[J]. 草业学报, 2015, 24(3):70-79 YANG Y T. Carbon and nitrogen metabolism patterns of Medicago ruthenica and its response to low temperature stress[D]. Hohhot: Inner Mongolia Agricultural University, 2022:105-106 杨艳婷. 扁蓿豆的碳氮代谢规律及其对低温胁迫的响应[D]. 呼和浩特:内蒙古农业大学, 2022:105-106 WANG M, LI L, JIA R, et al. Physiological characteristics and cold tolerance evaluation of 10 alfalfa varieties under low temperature stress[J]. Acta Prataculturae Sinica, 2024, 33(6):76-88 王敏, 李莉, 贾蓉, 等. 10种紫花苜蓿在低温胁迫下的生理特性及耐寒性评价[J]. 草业学报, 2024, 33(6):76-88 LIU Y, TIKUNOV Y, SCHOUTEN R E, et al. Anthocyanin biosynthesis and degradation mechanisms in solanaceous vegetables: A Review[J]. Frontiers in Chemistry, 2018, 6:52 LI Y T, ZHANG J, WANG S J, et al. Integrative transcriptomic and metabolomic analyses reveal the flavonoid biosynthesis of Pyrus hopeiensis flowers under cold stress[J]. Horticultural Plant Journal, 2023, 9(3):395-413 CHEN Y, CHEN L, SUN X M, et al. The mitogen-activated protein kinase kinase MKK2 positively regulates constitutive cold resistance in the potato[J]. Environmental and Experimental Botany, 2022, 194:104702)
Physiological Changes and Transcriptomic Analysis of Caucasian Clover under Low Temperature Stress
WU Qian1, ZHAO Ya-ru2, CAO Ke-fan1, MA Yi-ming1, SUO Yuan1, LI Shu-ning1, QI Hai-bo1, WANG Ming-jiu1
Affiliations
1 College of Grassland Science, Inner Mongolia Agricultural University/Key Laboratory of Grassland Resource of Ministry of Education, Hohhot, Inner Mongolia 010011, China; 2 Inner Mongolia Zhengshi Ecological Agriculture (Group) Co., Ltd., Hohhot, Inner Mongolia 010010, China
doi: 10.11733/j.issn.1007-0435.2026.07.006
文章导航
摘要
收起
本研究以耐寒性强的多年生牧草高加索三叶草(Trifolium ambiguum M. Bieb.)为材料,以20℃正常生长的幼苗为对照(CK),设置4℃低温胁迫2 h,6 h 和 12 h共3个处理,测定各时期高加索三叶草的生理指标,并对CK与胁迫12 h的植株进行转录组测序,解析其幼苗期在低温胁迫下的生理反应与适应机制。结果表明,4℃处理下,植株通过提高抗氧化酶(过氧化物酶、过氧化氢酶、超氧化物歧化酶)活性及渗透调节物质(可溶性蛋白、可溶性糖、游离脯氨酸)含量,有效清除活性氧,减轻了膜脂过氧化损伤,从而增强其抗寒能力。转录组分析显示,4℃处理12 h后共鉴定到15 428个差异表达基因,其中8909个基因上调,6519个基因下调。其中类黄酮生物合成通路与MAPK信号通路被显著激活:类黄酮通路上游关键基因查尔酮合成酶和查尔酮异构酶显著上调,促进了抗氧化物质(木犀草素、芹菜素、异黄酮)的合成;MAPK通路中MEKK1等基因的表达升高,表明低温信号通过磷酸化级联反应进行转导。其研究结果为解析高加索三叶草耐寒性提供了理论依据。
关键词
高加索三叶草
/
低温胁迫
/
生理指标
/
差异基因
Abstract
收起
This study investigated the physiological and molecular mechanisms underlying the cold tolerance of Caucasian clover (Trifolium ambiguum M. Bieb.), a perennial forage known for its strong hardiness. Seedlings grown at 20℃ served as the control (CK), and low-temperature stress (4℃) was applied for 2, 6, and 12 h. Physiological indicators were measured at each time point, and transcriptome sequencing was conducted on the control and 12 h-stressed plants. The results indicated that under 4℃, the plants enhanced their cold tolerance by significantly increasing the activities of antioxidant enzymes (peroxidase, catalase, and superoxide dismutase) and the levels of osmoregulatory substances (soluble protein, soluble sugar, and free proline). This coordinated response effectively scavenged reactive oxygen species and mitigated membrane lipid peroxidation, as evidenced by reduced malondialdehyde accumulation. Transcriptome analysis identified 15 428 differentially expressed genes (DEGs) after 12 h of cold stress, with 8909 up-regulated and 6519 down-regulated. Key pathways were significantly activated, including the flavonoid biosynthesis pathway, where the up-regulation of pivotal upstream genes (CHS and CHI) facilitated the synthesis of antioxidant compounds such as luteolin, apigenin, and isoflavones. Concurrently, the MAPK signaling pathway was induced, highlighted by the increased expression of genes like MEKK1, suggesting that cold signals are transduced via a phosphorylation cascade. These findings provide crucial insights into the molecular basis of cold adaptation in Caucasian clover.
WU Qian, ZHAO Ya-ru, CAO Ke-fan, MA Yi-ming, SUO Yuan, LI Shu-ning, QI Hai-bo, WANG Ming-jiu.
Physiological Changes and Transcriptomic Analysis of Caucasian Clover under Low Temperature Stress[J].
Acta Agrestia Sinica,
2026
, 34
(7)
: 2403
-2414
.
DOI: 10.11733/j.issn.1007-0435.2026.07.006
ZHENG Y F, CHEN J L, XIAO F, et al. Effects of different temperatures on photosynthetic characteristics and expression of photosynthesis-related genes in amaranth[J]. Journal of Plant Resources and Environment, 2022, 31(5):50-57 郑友峰, 陈家兰, 肖昉, 等 不同温度对苋菜光合特性及光合作用相关基因表达的影响[J]. 植物资源与环境学报, 2022, 31(5):50-57 DING Y L, SHI Y T, YANG S H. Advances and challenges in uncovering cold tolerance regulatory mechanisms in plants[J]. New phytologist, 2019, 222(4):1690-1704 XIONG H Y, DUAN R J, WANG R S. Advances in physiological basis and molecular mechanisms of cold acclimation affecting photosynthetic characteristics and plant type characteristics in overwintering crops[J]. Crops, 2017, (4):21-26 熊辉岩, 段瑞君, 王瑞生. 冷驯化影响越冬作物光合特性和株型特征的生理基础与分子机制的研究进展[J]. 作物杂志, 2017, (4):21-26 XIANG H T, ZHENG D F, HE N, et al. Research progress on physiological responses of plants to low temperature stress and mitigation effects of exogenous abscisic acid[J]. Acta Prataculturae Sinica, 2021, 30(1):208-219 项洪涛, 郑殿峰, 何宁, 等. 植物对低温胁迫的生理响应及外源脱落酸缓解胁迫效应的研究进展[J]. 草业学报, 2021, 30(1):208-219 HUANG L F, LONG Y Z, LI J Q, et al. Effects of low temperature stress on physiological characteristics of Coffea arabica L. seedlings[J]. Journal of Agricultural Science and Technology, 2023, 25(2):60-67 黄丽芳, 龙宇宙, 李金芹, 等. 低温胁迫对小粒种咖啡幼苗生理特性的影响[J]. 中国农业科技导报, 2023, 25(2):60-67 WEI X X, LI Y M, LI Y B, et al. Correlation between cold resistance and phloem sugar content and endogenous hormone accumulation in grape[J]. Acta Agriculturae Boreali-occidentalis Sinica, 2021, 30(3):386-394 韦霞霞, 李玉梅, 李彦彪, 等. 葡萄抗寒性与韧皮部糖分及内源激素积累的相关性[J]. 西北农业学报, 2021, 30(3):386-394 YANG H Y, LIU C Z. Effects of low temperature stress on cold resistance physiological indexes of four Euonymus plants[J]. Grassland and Turf, 2020, 40(2):92-98 杨航宇, 刘长仲. 低温胁迫对4种卫矛属植物抗寒生理指标的影响[J]. 草原与草坪, 2020, 40(2):92-98 TAN H Q, HUANG H T, TIE M M, et al. Transcriptome profiling of two asparagus bean (Vigna unguiculata subsp. sesquipedalis) cultivars differing in chilling tolerance under cold stress[J]. PLoS One, 2017, 11(3):e0151105 SHEAFFER C C, MARTEN G C. Kura clover forage yield, forage quality, and stand dynamics[J]. Canadian Journal of Plant Science, 1991, 71(4):1169-1172 TOWNSEND C E. Phenotypic diversity for agronmic charaaters and frequency of self-compatible plants in Trifolum ambiguum[J]. Canadian Journal of Plant Science, 1970, 50(3):331-338 SPEER G S, ALLINSON D W. Kura clover (Trifolium ambiguum): Legume for forage and soil conservation[J]. Economic Botany, 1985, 39(2):165-176 YIN X J. Study on adaptability and utilization value of Trifolium ambiguum M. Bieb. in Inner Mongolia[D]. Hohhot: Inner Mongolia Agricultural University, 2004:5-9 殷秀杰. 高加索三叶草在内蒙古地区适应性和利用价值研究[D]. 呼和浩特:内蒙古农业大学, 2004:5-9 MARSHALL A H, RASCLE C, ABBERTON M T, et al. Introgression as a route to improved drought tolerance in white clover (Trifolium repens L.)[J]. Journal of Agronomy and Crop Science, 2001, 187(1):11-18 NGUYEN T D, ANDREWS M, LEY T W S,, et al. Caucasian clover (Trifolium ambiguum) specific rhizobia persist in low and high fertility soils in the South Island of New Zealand[J]. New Zealand Journal of Agricultural Research, 2020, 63(3):332-340 PENG X, SHI F L, FU N N. Relationship between pigment, polyphenol contents and leaf color parameters in Trifolium lupinaster leaves[J]. Acta Agrestia Sinica, 2023, 31(7):2005-2011 彭新, 石凤翎, 付娜娜. 野火球叶片色素和多酚类物质含量与叶色参数的关系[J]. 草地学报, 2023, 31(7):2005-2011 ZHUANG D F, WANG C, BAO J H, et al. Antioxidant enzyme activities and related gene expressions of Rosa xanthina F. normalis under low temperature stress[J]. Journal of Northeast Forestry University, 2023, 51(4):89-92 庄得凤, 王成, 包金花, 等. 低温胁迫下单瓣黄刺玫抗氧化酶活性及相关基因表达[J]. 东北林业大学学报, 2023, 51(4):89-92 WEN R Y, LIU J X, LI S, et al. Effects of low temperature stress on physiological and biochemical characteristics of different quinoa seedlings[J]. Seed, 2019, 38(5):53-56 温日宇, 刘建霞, 李顺, 等. 低温胁迫对不同藜麦幼苗生理生化特性的影响[J].种子, 2019, 38(5):53-56 SUN Y L, LI J D, SUN B, et al. Response of main cold resistance physiological indexes of different alfalfa varieties to low temperature[J]. Journal of Shenyang Agricultural University, 2017, 48(5):591-596 孙予璐, 李建东, 孙备, 等. 不同品种紫花苜蓿主要抗寒生理指标对低温的响应[J]. 沈阳农业大学学报, 2017, 48(5):591-596 GAO T T, MAO P C, TIAN X X, et al. Evaluation of cold tolerance of 10 white clover varieties at seedling stage[J]. Pratacultural Science, 2014, 31(9):1724-1731 高婷婷, 毛培春, 田小霞, 等. 10个白三叶品种苗期耐寒性评价[J]. 草业科学, 2014, 31(9):1724-1731 WU Y X, CAI C Y, TANG S B, et al. Research progress on growth, development and molecular mechanisms of plants in response to low temperature[J]. Jiangsu Agricultural Sciences, 2023, 51(19):1-9 吴宇欣, 蔡昌杨, 唐诗蓓, 等. 植物响应低温的生长发育及分子机制研究进展[J]. 江苏农业科学, 2023, 51(19):1-9 SU L W, LI S, MA S Y, et al. Establishment of a comprehensive evaluation method for grape cold resistance[J]. Acta Prataculturae Sinica, 2015, 24(3):70-79 苏李维, 李胜, 马绍英, 等. 葡萄抗寒性综合评价方法的建立[J]. 草业学报, 2015, 24(3):70-79 YANG Y T. Carbon and nitrogen metabolism patterns of Medicago ruthenica and its response to low temperature stress[D]. Hohhot: Inner Mongolia Agricultural University, 2022:105-106 杨艳婷. 扁蓿豆的碳氮代谢规律及其对低温胁迫的响应[D]. 呼和浩特:内蒙古农业大学, 2022:105-106 WANG M, LI L, JIA R, et al. Physiological characteristics and cold tolerance evaluation of 10 alfalfa varieties under low temperature stress[J]. Acta Prataculturae Sinica, 2024, 33(6):76-88 王敏, 李莉, 贾蓉, 等. 10种紫花苜蓿在低温胁迫下的生理特性及耐寒性评价[J]. 草业学报, 2024, 33(6):76-88 LIU Y, TIKUNOV Y, SCHOUTEN R E, et al. Anthocyanin biosynthesis and degradation mechanisms in solanaceous vegetables: A Review[J]. Frontiers in Chemistry, 2018, 6:52 LI Y T, ZHANG J, WANG S J, et al. Integrative transcriptomic and metabolomic analyses reveal the flavonoid biosynthesis of Pyrus hopeiensis flowers under cold stress[J]. Horticultural Plant Journal, 2023, 9(3):395-413 CHEN Y, CHEN L, SUN X M, et al. The mitogen-activated protein kinase kinase MKK2 positively regulates constitutive cold resistance in the potato[J]. Environmental and Experimental Botany, 2022, 194:104702