Article(id=1312006164238397652, tenantId=1146029695717560320, journalId=1311385948879159419, issueId=1312006144101540779, articleNumber=null, orderNo=null, doi=10.13430/j.cnki.jpgr.20250919003, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1758211200000, receivedDateStr=2025-09-19, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1790736245450, onlineDateStr=2026-09-30, pubDate=1784131200000, pubDateStr=2026-07-16, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1790736245450, onlineIssueDateStr=2026-09-30, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1790736245450, creator=13701087609, updateTime=1790736245450, updator=13701087609, issue=Issue{id=1312006144101540779, tenantId=1146029695717560320, journalId=1311385948879159419, year='2026', volume='27', issue='7', pageStart='1311', pageEnd='1484', issueExtLink='null', onlineDate='null', pubDate='1784131200000', pubDateStr='2026-07-16', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1790736240648, creator='13701087609', updateTime=1790739525097, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1312019920167203355, tenantId=1146029695717560320, journalId=1311385948879159419, issueId=1312006144101540779, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1312019920167203356, tenantId=1146029695717560320, journalId=1311385948879159419, issueId=1312006144101540779, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1377, endPage=1387, ext={EN=ArticleExt(id=1312006166008393941, articleId=1312006164238397652, tenantId=1146029695717560320, journalId=1311385948879159419, language=EN, title=Research Progress of Mining Disease-resistant Genes in Elytrigia and Their Breeding Applications, columnId=1312006144902652845, journalTitle=Journal of Plant Genetic Resources, columnName=REVIEW, runingTitle=null, highlight=null, articleAbstract=

The genus Elytrigia (Thinopyrum) exhibit outstanding disease resistance and serve as an important member of the tertiary gene pool of wheat. The exploration and application of disease-resistant genes from Elytrigia have provided critical support for the wheat improvement in disease resistance. As early as the 1920s, reports on distant hybridization between wheat and Elytrigia emerged. Over the past century, numerous disease-resistant genes have been identified in Elytrigia species and multiple disease-resistant germplasm have been developed by distant hybridization and molecular chromosome engineering and breeding of high-yielding and disease-resistant cultivars such as Xiaoyan 6, which have made significant contributions to wheat breeding and food security. This article systematically reviews the general of Elytrigia genus species, the research progress on genes resistant to common fungal diseases such as stripe rust, leaf rust, stem rust, powdery mildew, fusarium head blight, and crown rot, and summarizes advances in creating novel germplasm and breeding applications of disease-resistant genes through distant hybridization and molecular chromosome engineering. Furthermore, it discusses challenges in the exploration and application of disease-resistant genes from Elytrigia, and prospects future research directions. This review is structured into the aforementioned four sections, aiming to provide a scientific basis for in-depth analysis of the mechanisms of exogenous disease-resistant genes and enhance their utilization efficiency in wheat breeding programs.

, authors=Xinyue FAN1, Ruotong LI1, Xinyue HU1, Qi ZHOU1, Tongfei QI1, Yao LEI1, Yunxiang WANG3, Chunyan TONG1, 2, Xingfeng LI1, 2, Yinguang BAO1, 2, Guotang YANG1, 2, authorsList=Xinyue FAN, Ruotong LI, Xinyue HU, Qi ZHOU, Tongfei QI, Yao LEI, Yunxiang WANG, Chunyan TONG, Xingfeng LI, Yinguang BAO, Guotang YANG, authorCompany=null, correspAuthors=Guotang YANG, 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=1312006166637539542, articleId=1312006164238397652, tenantId=1146029695717560320, journalId=1311385948879159419, language=CN, title=偃麦草属抗病基因挖掘及育种应用进展, columnId=1312006145041064879, journalTitle=植物遗传资源学报, columnName=综述, runingTitle=null, highlight=null, articleAbstract=

偃麦草属(Elytrigia,异名为Thinopyrum)植物抗病性能突出,是小麦三级基因库中的重要成员,其抗病基因的挖掘与应用为小麦抗病改良提供了关键支撑。早在20世纪20年代,就有小麦与偃麦草远缘杂交的报道。历经100余年,在偃麦草属中鉴定到多个抗病基因,通过远缘杂交与分子染色体工程创制出多份抗病种质并培育出以小偃6号为代表的高产、抗病新品种,为小麦育种及粮食安全做出了突出贡献。本文系统综述了偃麦草属植物概况,以及抗条锈病、叶锈病、秆锈病、白粉病、赤霉病和茎基腐病等常见真菌病害基因挖掘的研究进展,总结了通过远缘杂交、分子染色体工程等手段创制新种质和抗病基因的育种应用进展。另外,讨论了偃麦草属抗病基因挖掘及应用过程中存在的问题,并展望了未来发展方向。本文旨在为深入解析外源抗病基因的作用机制和在小麦育种过程中提高其利用效率提供科学依据。

, authors=范馨月1, 李若彤1, 胡心悦1, 周琦1, 亓同飞1, 雷垚1, 王韵翔3, 佟春艳1, 2, 李兴锋1, 2, 鲍印广1, 2, 杨国堂1, 2, authorsList=范馨月, 李若彤, 胡心悦, 周琦, 亓同飞, 雷垚, 王韵翔, 佟春艳, 李兴锋, 鲍印广, 杨国堂, authorCompany=null, correspAuthors=杨国堂, authorNote=

研究方向为小麦种质创新,E-mail:

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杨国堂,研究方向为小偃麦分子染色体工程,E-mail:
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研究方向为小麦种质创新,E-mail:

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研究方向为小麦种质创新,E-mail:

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Nature Genetics, 2023, 55(6):914-920, articleTitle=An unusual tandem kinase fusion protein confers leaf rust resistance in wheat, refAbstract=null), Reference(id=1312006188645052811, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, doi=null, pmid=null, pmcid=null, year=2023, volume=14, issue=1, pageStart=4353, pageEnd=null, url=null, language=null, rfNumber=77, rfOrder=95, authorNames=Ni F, Zheng Y Y, Liu X K, Yu Y, Zhang G Q, Epstein L, Mao X, Wu J Z, Yuan C L, Lv B, Yu H X, Li J L, Zhao Q, Yang Q Y, Liu J J, Qi J, Fu D L, Wu J J, journalName=Nature Communications, refType=null, unstructuredReference=Ni F, Zheng Y Y, Liu X K, Yu Y, Zhang G Q, Epstein L, Mao X, Wu J Z, Yuan C L, Lv B, Yu H X, Li J L, Zhao Q, Yang Q Y, Liu J J, Qi J, Fu D L, Wu J J. Sequencing trait-associated mutations to clone wheat rust-resistance gene YrNAM. Nature Communications, 2023, 14(1):4353, articleTitle=Sequencing trait-associated mutations to clone wheat rust-resistance gene YrNAM, refAbstract=null), Reference(id=1312006188716355980, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, doi=null, pmid=null, pmcid=null, year=2023, volume=136, issue=7, pageStart=148, pageEnd=null, url=null, language=null, rfNumber=78, rfOrder=96, authorNames=Wan W T, Zhao R H, Chen T T, Wang L, Zhang X, Li H F, Wang X E, Bie T D, journalName=Theoretical and Applied Genetics, refType=null, unstructuredReference=Wan W T, Zhao R H, Chen T T, Wang L, Zhang X, Li H F, Wang X E, Bie T D. Rapid development of wheat-Dasypyrum villosum compensating translocations resistant to powdery mildew using a triple marker strategy conducted on a large ph1b-induced population. 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Application of dwarf-male-sterile wheat in breeding program for Fusarium head blight resistance in Southern Yellow-Huai Rivers Valley Winter Wheat Region. Acta Agronomica Sinica, 2022, 48(11): 2683-2690, articleTitle=Application of dwarf-male-sterile wheat in breeding program for Fusarium head blight resistance in Southern Yellow-Huai Rivers Valley Winter Wheat Region, refAbstract=null)], funds=null, companyList=[AuthorCompany(id=1312006166981472471, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, xref=1, ext=[AuthorCompanyExt(id=1312006166989861080, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006166981472471, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1Agronomy College, Shandong Agricultural University, Tai′an 271018), AuthorCompanyExt(id=1312006166998249689, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006166981472471, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1山东农业大学农学院,泰安 271018)]), AuthorCompany(id=1312006167073747162, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, xref=2, ext=[AuthorCompanyExt(id=1312006167082135771, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006167073747162, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2State Key Laboratory of Wheat Improvement, Tai′an 271018, Shandong), AuthorCompanyExt(id=1312006167090524380, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006167073747162, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2小麦育种全国重点实验室,山东 泰安 271018)]), AuthorCompany(id=1312006167170216157, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, xref=3, ext=[AuthorCompanyExt(id=1312006167182799070, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006167170216157, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3College of Future Technology (Qilu College), Shandong Agricultural University, Tai′an 271018), AuthorCompanyExt(id=1312006167191187679, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, companyId=1312006167170216157, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3山东农业大学未来技术学院(齐鲁学堂),泰安 271018)])], figs=[ArticleFig(id=1312006172689920298, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, language=EN, label=Table 1, caption=

The major fungal disease resistance genes in Elytrigia

, figureFileSmall=null, figureFileBig=null, tableContent=

基因

Genes

来源

Sources

抗病特性

Resistance characteristics

染色体定位

Chromosomal localizations

参考文献

References

Yr50中间偃麦草抗条锈病4B[19]
YrL693中间偃麦草抗条锈病1B[20]
WCBP1中间偃麦草抗条锈病1B[21]
YrCH-1BS中间偃麦草抗条锈病3Ai[22]
YrT14中间偃麦草抗条锈病7J或7Js[23]
Yr69十倍体长穗偃麦草抗条锈病2A[24]
YrTp1十倍体长穗偃麦草抗条锈病2B[25]
YrTp2十倍体长穗偃麦草抗条锈病7B[25]
YrXY十倍体长穗偃麦草抗条锈病4B[26]
Yr4EL四倍体长穗偃麦草抗条锈病4E[27]
Lr19十倍体长穗偃麦草抗叶锈病7Ae#1[29-30]
Lr24十倍体长穗偃麦草抗叶锈病3Ae#1[31-33]
Lr29十倍体长穗偃麦草抗叶锈病7Ae#1[34-35]
Lr5Ag十倍体长穗偃麦草抗叶锈病5Ag[37]
Lr38中间偃麦草抗叶锈病7Ai#2[36]
Sr24十倍体长穗偃麦草抗秆锈病3Ae#1[31-33]
Sr25十倍体长穗偃麦草抗秆锈病7Ae#1[29]
Sr26十倍体长穗偃麦草抗秆锈病6Ae#1[10, 38]
Sr43十倍体长穗偃麦草抗秆锈病7el2[11, 39-43]
Sr61十倍体长穗偃麦草抗秆锈病6Ae#3[10, 44]
Sr44中间偃麦草抗秆锈病7Ai-1[36, 45]
Sr68灯芯偃麦草抗秆锈病4E[46]
Pm51十倍体长穗偃麦草抗白粉病2B[48]
Pm5Ag十倍体长穗偃麦草抗白粉病5Ag[37]
Pm40中间偃麦草抗白粉病7B[49]
Pm43中间偃麦草抗白粉病2D[50]
pmCH83中间偃麦草抗白粉病4B[51]
PmCH7124中间偃麦草抗白粉病2B[52]
pmCH89中间偃麦草抗白粉病4B[53]
PmL692中间偃麦草抗白粉病2B[54]
Fhb7十倍体长穗偃麦草抗赤霉病、抗茎基腐病7el2[12]
TTE7E-Fhb7四倍体长穗偃麦草抗赤霉病7E[59]
Fhb7The2二倍体长穗偃麦草抗赤霉病7E[57]
Fhb7-7EL二倍体长穗偃麦草抗赤霉病、抗茎基腐病7E[58, 61]
QFhb.nwafu-5B十倍体长穗偃麦草抗赤霉病5B[60]
QFhb.nwafu-7A十倍体长穗偃麦草抗赤霉病7A[60]
), ArticleFig(id=1312006172786389291, tenantId=1146029695717560320, journalId=1311385948879159419, articleId=1312006164238397652, language=CN, label=表1, caption=

偃麦草属主要的抗真菌病害基因

, figureFileSmall=null, figureFileBig=null, tableContent=

基因

Genes

来源

Sources

抗病特性

Resistance characteristics

染色体定位

Chromosomal localizations

参考文献

References

Yr50中间偃麦草抗条锈病4B[19]
YrL693中间偃麦草抗条锈病1B[20]
WCBP1中间偃麦草抗条锈病1B[21]
YrCH-1BS中间偃麦草抗条锈病3Ai[22]
YrT14中间偃麦草抗条锈病7J或7Js[23]
Yr69十倍体长穗偃麦草抗条锈病2A[24]
YrTp1十倍体长穗偃麦草抗条锈病2B[25]
YrTp2十倍体长穗偃麦草抗条锈病7B[25]
YrXY十倍体长穗偃麦草抗条锈病4B[26]
Yr4EL四倍体长穗偃麦草抗条锈病4E[27]
Lr19十倍体长穗偃麦草抗叶锈病7Ae#1[29-30]
Lr24十倍体长穗偃麦草抗叶锈病3Ae#1[31-33]
Lr29十倍体长穗偃麦草抗叶锈病7Ae#1[34-35]
Lr5Ag十倍体长穗偃麦草抗叶锈病5Ag[37]
Lr38中间偃麦草抗叶锈病7Ai#2[36]
Sr24十倍体长穗偃麦草抗秆锈病3Ae#1[31-33]
Sr25十倍体长穗偃麦草抗秆锈病7Ae#1[29]
Sr26十倍体长穗偃麦草抗秆锈病6Ae#1[10, 38]
Sr43十倍体长穗偃麦草抗秆锈病7el2[11, 39-43]
Sr61十倍体长穗偃麦草抗秆锈病6Ae#3[10, 44]
Sr44中间偃麦草抗秆锈病7Ai-1[36, 45]
Sr68灯芯偃麦草抗秆锈病4E[46]
Pm51十倍体长穗偃麦草抗白粉病2B[48]
Pm5Ag十倍体长穗偃麦草抗白粉病5Ag[37]
Pm40中间偃麦草抗白粉病7B[49]
Pm43中间偃麦草抗白粉病2D[50]
pmCH83中间偃麦草抗白粉病4B[51]
PmCH7124中间偃麦草抗白粉病2B[52]
pmCH89中间偃麦草抗白粉病4B[53]
PmL692中间偃麦草抗白粉病2B[54]
Fhb7十倍体长穗偃麦草抗赤霉病、抗茎基腐病7el2[12]
TTE7E-Fhb7四倍体长穗偃麦草抗赤霉病7E[59]
Fhb7The2二倍体长穗偃麦草抗赤霉病7E[57]
Fhb7-7EL二倍体长穗偃麦草抗赤霉病、抗茎基腐病7E[58, 61]
QFhb.nwafu-5B十倍体长穗偃麦草抗赤霉病5B[60]
QFhb.nwafu-7A十倍体长穗偃麦草抗赤霉病7A[60]
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偃麦草属抗病基因挖掘及育种应用进展
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范馨月 1 , 李若彤 1 , 胡心悦 1 , 周琦 1 , 亓同飞 1 , 雷垚 1 , 王韵翔 3 , 佟春艳 1, 2 , 李兴锋 1, 2 , 鲍印广 1, 2 , 杨国堂 1, 2
植物遗传资源学报 | 综述 2026,27(7): 1377-1387
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植物遗传资源学报 |综述 2026 , 27 (7) : 1377 -1387
偃麦草属抗病基因挖掘及育种应用进展
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范馨月1 , 李若彤1, 胡心悦1, 周琦1, 亓同飞1, 雷垚1, 王韵翔3, 佟春艳1, 2, 李兴锋1, 2, 鲍印广1, 2, 杨国堂1, 2
作者信息
  • 1山东农业大学农学院,泰安 271018
  • 2小麦育种全国重点实验室,山东 泰安 271018
  • 3山东农业大学未来技术学院(齐鲁学堂),泰安 271018
通讯作者:
杨国堂,研究方向为小偃麦分子染色体工程,E-mail:
作者简介:

研究方向为小麦种质创新,E-mail:

Research Progress of Mining Disease-resistant Genes in Elytrigia and Their Breeding Applications
Xinyue FAN1 , Ruotong LI1, Xinyue HU1, Qi ZHOU1, Tongfei QI1, Yao LEI1, Yunxiang WANG3, Chunyan TONG1, 2, Xingfeng LI1, 2, Yinguang BAO1, 2, Guotang YANG1, 2
Affiliations
  • 1Agronomy College, Shandong Agricultural University, Tai′an 271018
  • 2State Key Laboratory of Wheat Improvement, Tai′an 271018, Shandong
  • 3College of Future Technology (Qilu College), Shandong Agricultural University, Tai′an 271018
出版时间: 2026-07-16 doi: 10.13430/j.cnki.jpgr.20250919003
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偃麦草属(Elytrigia,异名为Thinopyrum)植物抗病性能突出,是小麦三级基因库中的重要成员,其抗病基因的挖掘与应用为小麦抗病改良提供了关键支撑。早在20世纪20年代,就有小麦与偃麦草远缘杂交的报道。历经100余年,在偃麦草属中鉴定到多个抗病基因,通过远缘杂交与分子染色体工程创制出多份抗病种质并培育出以小偃6号为代表的高产、抗病新品种,为小麦育种及粮食安全做出了突出贡献。本文系统综述了偃麦草属植物概况,以及抗条锈病、叶锈病、秆锈病、白粉病、赤霉病和茎基腐病等常见真菌病害基因挖掘的研究进展,总结了通过远缘杂交、分子染色体工程等手段创制新种质和抗病基因的育种应用进展。另外,讨论了偃麦草属抗病基因挖掘及应用过程中存在的问题,并展望了未来发展方向。本文旨在为深入解析外源抗病基因的作用机制和在小麦育种过程中提高其利用效率提供科学依据。

普通小麦  /  偃麦草属  /  远缘杂交  /  抗病基因  /  种质创新  /  育种利用

The genus Elytrigia (Thinopyrum) exhibit outstanding disease resistance and serve as an important member of the tertiary gene pool of wheat. The exploration and application of disease-resistant genes from Elytrigia have provided critical support for the wheat improvement in disease resistance. As early as the 1920s, reports on distant hybridization between wheat and Elytrigia emerged. Over the past century, numerous disease-resistant genes have been identified in Elytrigia species and multiple disease-resistant germplasm have been developed by distant hybridization and molecular chromosome engineering and breeding of high-yielding and disease-resistant cultivars such as Xiaoyan 6, which have made significant contributions to wheat breeding and food security. This article systematically reviews the general of Elytrigia genus species, the research progress on genes resistant to common fungal diseases such as stripe rust, leaf rust, stem rust, powdery mildew, fusarium head blight, and crown rot, and summarizes advances in creating novel germplasm and breeding applications of disease-resistant genes through distant hybridization and molecular chromosome engineering. Furthermore, it discusses challenges in the exploration and application of disease-resistant genes from Elytrigia, and prospects future research directions. This review is structured into the aforementioned four sections, aiming to provide a scientific basis for in-depth analysis of the mechanisms of exogenous disease-resistant genes and enhance their utilization efficiency in wheat breeding programs.

common wheat  /  Elytrigia  /  distant hybridization  /  disease-resistant gene  /  germplasm innovation  /  breeding utilization
范馨月, 李若彤, 胡心悦, 周琦, 亓同飞, 雷垚, 王韵翔, 佟春艳, 李兴锋, 鲍印广, 杨国堂. 偃麦草属抗病基因挖掘及育种应用进展. 植物遗传资源学报, 2026 , 27 (7) : 1377 -1387 . DOI: 10.13430/j.cnki.jpgr.20250919003
Xinyue FAN, Ruotong LI, Xinyue HU, Qi ZHOU, Tongfei QI, Yao LEI, Yunxiang WANG, Chunyan TONG, Xingfeng LI, Yinguang BAO, Guotang YANG. Research Progress of Mining Disease-resistant Genes in Elytrigia and Their Breeding Applications[J]. Journal of Plant Genetic Resources, 2026 , 27 (7) : 1377 -1387 . DOI: 10.13430/j.cnki.jpgr.20250919003
普通小麦(Triticum aestivum L., 2n = 6x = 42)包含A、B和D三套亚基因组,是由小麦属(Triticum)和山羊草属(Aegilops)植物经两次自然杂交形成的异源六倍体[1]。我国是小麦产能与消费第一大国,其中,2024年我国小麦总产量达到1.4亿t,约占全球17%,而小麦年消费量为1.2亿t,约占全球19%(https://knoema.com)。由此可见,培育高产、稳产的小麦品种对保障国家粮食安全、保证国民经济可持续性发展具有重要意义。近年来,随着科技创新,小麦的育种水平不断提升。但长期的选择压力导致其遗传基础逐渐狭窄,对病虫害的抵抗能力不断下降。Savary等[2]的研究表明,在全世界已报道的31种小麦病虫害中,叶锈病(Leaf rust)、赤霉病(Fusarium head blight)、叶枯病(Tritici blotch)、条锈病(Stripe rust)、蠕孢叶枯病(Spot blotch)、褐斑病(Tan spot)、蚜虫(Aphids)、白粉病(Powdery mildew)等8种对小麦生产造成主要威胁。在我国,小麦锈病、白粉病、赤霉病等真菌病害频发,近年来,茎基腐病发病程度也呈逐年上升趋势。因此,防治这些小麦真菌病害对于维护小麦稳产十分关键。小麦野生近缘种中含有丰富的抗病、耐逆基因。通过远缘杂交和染色体工程的方法,将野生近缘种中的优异基因导入小麦,能够极大地拓宽小麦的遗传基础,为小麦生产提供有力保障。野生近缘种在提升栽培种抗逆特性方面具有巨大育种潜力,被中国科协列为2025年十大前沿科学问题[3]。
偃麦草属(Elytrigia,异名为Thinopyrum)植物作为小麦重要的野生资源之一,具有抗病、耐逆、高籽粒蛋白质含量等优良性状且易与小麦杂交,被广泛应用于小麦育种[4]。在抗非生物胁迫方面,十倍体长穗偃麦草(E. elongata,异名为Th. ponticum)的耐盐特性应用于小麦育种,培育出山融3号、小偃60等耐盐新品种[5-6]。在优质育种方面,将十倍体长穗偃麦草与中间偃麦草(E. intermedia,异名为Th. intermedium)的优质基因转入小麦,培育出小偃54、小冰麦33、高优503等优质新品种[7-9]。在抗生物胁迫方面,偃麦草属抗病基因的挖掘与应用方面取得明显进展。目前,已正式命名17个抗病基因,其中Sr26、Sr43、Sr61、Fhb7等4个基因已被克隆,Fhb7的赤霉病解毒机制也已解析[10-12]。尽管如此,在抗病基因挖掘方面仍面临诸多挑战,如多数抗病基因尚未克隆,其分子机制尚未阐明;多数携带抗性基因的染色体片段存在连锁累赘,导致育种利用效率不高。本文综述了偃麦草属中主要抗真菌病害基因挖掘及其在小麦种质创新和育种应用中的最新研究进展,以期为小麦远缘杂交和抗病育种工作者提供参考。
偃麦草属植物是禾本科小麦族多年生草本植物,1933年Nevski利用系统分类法将偃麦草属从传统的冰草属(Agropyron)中分离出来,独立成属[13]。而后,我国植物分类学家耿以礼和郭本兆沿用了该分类方法[14-15]。偃麦草属植物在全球有40余种,其中,长穗偃麦草、中间偃麦草、毛偃麦草(E. trichophora)、硬叶偃麦草(E. smithii)和脆轴偃麦草(E. juncea)等被广泛应用[16]。中间偃麦草是异源六倍体,而长穗偃麦草因其倍性不同又划分为二倍体、四倍体和十倍体3类。中间偃麦草为常异花授粉植物,具有短而匍匐的地下茎,茎秆较软,一般株高1.2~1.8 m,穗长20~40 cm,每穗小穗数20~30个,每小穗花数5~9个,千粒重4~6 g,结实率低。中间偃麦草对锈病、白粉病、叶枯病等生物胁迫以及寒冷、盐渍及干旱等非生物胁迫均具有很强的抗性[16]。十倍体长穗偃麦草为常异花授粉植物,根系强大,再生能力强,分蘖多,茎秆坚硬;单株成穗可达100个以上,一般株高1.5~2.0 m,穗长40~50 cm,每穗小穗数30个左右,每小穗花数5~10个,每穗结实粒数100粒左右,千粒重4 g左右。十倍体长穗偃麦草对锈病、白粉病、赤霉病等病害具有很强的抗性[16],对低温、抗旱、盐碱等非生物胁迫也具有很好的耐受性。
截至2025年底,偃麦草属的二倍体长穗偃麦草(E. elongata,异名为Th. elongatum, 2n = 2x = 14, EE)和中间偃麦草(2n = 6x = 42, StStVVJJ)的基因组已公布[12, 17]。研究显示,E基因组为4.78 Gb,重复序列占比为81.29%,注释了44747个高可信度基因,与小麦A、B和D基因组均具有很好的共线性[12]。中间偃麦草基因组大小为10.89 Gb,重复序列占比为77.15%,预测了133578个编码蛋白基因[17]。中间偃麦草共包含3个亚基因组,分别是来源于拟鹅观草(Pseudoroegneria libanotica,2n = 2x = 14, StSt)的St基因组、簇毛麦的V基因组以及与山羊草属D和S基因组相似的J基因组。此外,十倍体长穗偃麦草和中间偃麦草的祖先种拟鹅观草的基因组也已公布。研究显示,St基因组为2.99 Gb,重复序列占比为71.62%,注释了46369个编码蛋白基因[18]。从已公布的基因组数据得知,偃麦草属植物包含的亚基因组多且复杂,基因数目多,可为小麦遗传改良提供丰富的基因资源。
小麦条锈病是由条锈菌(Puccinia striiformis f. sp. tritici)引起的一种真菌病害。该病害在低温潮湿地区流行,严重影响小麦的产量。目前,已报道的来源于中间偃麦草的抗条锈病基因包括1个正式命名的基因Yr50,以及4个临时命名的基因YrCH-1BS、YrT14、YrL693、WCBP1[19-23]。其中,Yr50在小偃麦渐渗系CH223中定位于4B染色体的长臂,并介于Xbarc1096与Xwmc47两个SSR标记之间[19]。YrL693在抗病系L693中定位于1B染色体近着丝粒区,位于Xgwm273-1B和Xgwm187-1B两个标记之间[20]。WCBP1发现于抗病系L693,通过抑制消减杂交(SSH, suppression subtractive hybridization)技术获得,推测是YrL693的一个候选基因[21]。易位系CH4131和CH4132的易位染色体组成为T3Ai-1BS•1BL,成株期高抗条锈病,推测其携带的抗条锈病基因YrCH-1BS可能来源于中间偃麦草3Ai染色体[22]。YrT14来源于小麦与中间偃麦草异附加系TAI-14附加的中间偃麦草染色体7J(或7Js),物理区间为636.7 Mb~724.8 Mb[23]。同样,在十倍体长穗偃麦草中也正式命名了1个抗条锈病基因Yr69和3个临时命名基因YrTp1、YrTp2和YrXY[24-27]。其中,Yr69在小偃麦渐渗系CH7086中定位于2A染色体短臂末端,介于X2AS33和Xmag3807标记间[24]。显性基因YrTp1和隐性基因YrTp2均发现于小麦与长穗偃麦草远缘杂交后代A-3,YrTp1定位于2B染色体短臂,介于WMC477和Xgwm374两个标记之间;YrTp2定位于7B染色体短臂,位于WMC364和Xgwm333两个标记之间[25]。YrXY在小麦品种小偃6号中定位于小麦4B染色体,与标记Xgwm107的遗传距离为7.08 cM[26]。此外,普通小麦与四倍体长穗偃麦草(E. elongata,异名为Th. elongatum, 2n = 4x = 28, EEEE)4E(4D)代换系K16-730-3中还有1个临时命名的抗条锈病基因Yr4EL,该基因定位于4E染色体长臂[27]。
小麦叶锈病是由叶锈菌(Puccinia recondita f. sp. tritici)引起的一种真菌病害,严重威胁小麦的生产安全。据统计,叶锈病平均每年约造成15%的小麦产量损失,严重时可达50%[28]。叶锈菌适宜在白天温暖、晚上潮湿的环境中生长[1]。由于叶锈菌适应能力强,叶锈病的流行范围比条锈病和秆锈病的范围大,加之近年来气候变暖,叶锈病的流行区域呈现逐年扩大的趋势。目前,在长穗偃麦草中正式命名了Lr19、Lr24和Lr29共3个抗叶锈病基因,在中间偃麦草中正式命名了1个抗叶锈病基因Lr38[29-36]。其中,Lr19发现于小麦与长穗偃麦草易位系Agatha,位于7Ae#1染色体长臂[29],Xu等[30]进一步将其定位于长臂末端标记XsdauK3734和XsdauK2839之间。Lr24定位于小麦与长穗偃麦草易位系Agent的3Ae#1染色体长臂[31-33]。Lr29发现于7D/7Ae#1易位系,位于7Ae#1染色体短臂[34-35]。Lr38定位于小麦与中间偃麦草易位系W49的7Ai#2染色体长臂[36]。此外,在小麦与长穗偃麦草5Ag(5D)代换系的5Ag染色体中还临时命名了1个抗叶锈病基因Lr5Ag[37]。
小麦秆锈病是由秆锈菌(Puccinia graminis f. sp. tritici)引起的一种真菌病害,严重影响小麦产量及品质。目前,在长穗偃麦草中正式命名了Sr24、Sr25、Sr26、Sr43和Sr61共5个抗秆锈病基因,在中间偃麦草中正式命名了1个抗秆锈病基因Sr44,在灯芯偃麦草(E. junceum,异名为Th. junceum)中正式命名了1个抗秆锈病基因Sr68。Sr24发现于小麦与长穗偃麦草易位系Agent,位于3Ae#1染色体长臂[31-33]。Sr25发现于小麦与长穗偃麦草易位系Agatha,位于7Ae#1染色体长臂[29]。Sr26最初发现于小麦与长穗偃麦草杂交后代P.W.357-5,位于6Ae#1染色体长臂[38]。Sr43在小麦与长穗偃麦草易位系KS10-2和KS24-2中定位于7el2染色体长臂[39-43]。Sr61发现于小麦与长穗偃麦草代换系W3757,位于6Ae#3染色体长臂[44]。Sr44最初发现于小麦与中间偃麦草异附加系TAF2d,位于7Ai-1染色体短臂近末端[36, 45]。Sr68定位于小麦与灯芯偃麦草异附加系HD3505的4E染色体长臂[46]。Sr24与Lr24紧密连锁,Sr25与Lr19连锁,并且Sr25、Sr26、Sr43、Sr44、Sr61和Sr68对Ug99生理小种均具有抗性(https://rusttracker.cimmyt.org/?page_id=6745)。其中,Sr26、Sr43和Sr61已被正式克隆。Sr26和Sr61分别编码两个不同的NLR蛋白[10],Sr43编码一个含有未知结构域的蛋白激酶[11]。
小麦白粉病是由白粉菌(Blumeria graminis f. sp. tritici)引起的一种真菌病害,平均导致小麦5%~40%的产量损失,严重时高达62%[47]。目前,在长穗偃麦草中正式命名了1个抗白粉病基因Pm51,在中间偃麦草中正式命名了2个抗白粉病基因Pm40和Pm43。其中,Pm51在小麦与长穗偃麦草渐渗系CH7086中定位于2B染色体长臂,介于标记Xwmc332和BQ246670之间[48]。Pm40发现于小麦与中间偃麦草渐渗系GRY19,定位于7B染色体的短臂,与标记Xgwm297和Xwmc335共分离[49]。Pm43定位于小麦与中间偃麦草渐渗系CH5025的2D染色体长臂,介于标记Xwmc41和Xbarc11之间[50]。此外,在长穗偃麦草中临时命名了1个抗白粉病基因,在中间偃麦草中临时命名了4个抗白粉病基因,包括发现于小麦与长穗偃麦草5Ag(5D)代换系PY5814,定位于长穗偃麦草5Ag染色体上的Pm5Ag[37];发现于小麦与中间偃麦草渐渗系CH09W83,并定位于4B染色体长臂标记Xgwm251和Xwmc652之间的pmCH83[51];定位于渐渗系CH7124的2B染色体长臂标记Xgwm501和Xbarc101之间的PmCH7124[52];发现于渐渗系CH09W89,并定位于4B染色体长臂标记Xwmc310和Xwmc125之间的pmCH89[53]和定位于渐渗系L692的2B染色体短臂末端标记Xwmc314和BE443737之间的PmL692[54]。
小麦赤霉病是由禾谷镰刀菌(Fusarium graminearum)等多种镰刀菌引起的小麦穗部病害,严重威胁全球小麦产业发展。目前,在长穗偃麦草中仅正式命名了1个抗赤霉病基因Fhb7。该基因最初发现于小麦与长穗偃麦草代换系K2620,并定位于7el2染色体长臂[55]。Guo等[56]进一步将其精细定位于XsdauK66和Xcfa2240两个标记之间。Wang等[12]通过图位克隆,发现Fhb7编码一个谷胱甘肽巯基转移酶,该酶通过破坏关键制毒环氧基团实现呕吐毒素的降解。在二倍体长穗偃麦草7E染色体上,同源克隆了Fhb7的等位基因Fhb7The2[57];此外还临时命名1个抗赤霉病基因Fhb7-7EL[58]。在四倍体长穗偃麦草中,同样发现7E染色体携带抗赤霉病基因,并同源克隆出Fhb7的等位基因TTE7E-Fhb7[59]。此外,在小麦与长穗偃麦草远缘杂交后代品种西农511中,定位了2个抗赤霉病位点QFhb.nwafu-5B和QFhb.nwafu-7A,推测这2个抗性位点可能来源于长穗偃麦草基因组[60]。
小麦茎基腐病是由假禾谷镰孢菌(Fusarium pseudograminearum)、禾谷镰孢菌等引起的全球性土传病害。它在茎基部引发、全生育期均可产生危害。近年来在黄淮海等小麦主产区的发病呈范围扩大、危害程度加重的趋势,严重威胁小麦稳产。目前,偃麦草属中未有正式命名的抗茎基腐病基因,但存在相关抗性位点的报道。其中,Wang等[12]发现Fhb7不仅具有赤霉病抗性,也具有茎基腐病抗性。Ceoloni等[61]发现二倍体长穗偃麦草Fhb7-7EL位点同样具有茎基腐病抗性。表1总结了36个基因的来源、抗病特性和在偃麦草和小麦染色体的定位情况,为其进一步应用提供指导。
自20世纪20年代,国内外开始有通过远缘杂交和染色体工程技术创制小偃麦的报道[62]。其中,黑龙江省农业科学院利用中间偃麦草与普通小麦杂交,育成了一批“中”字系列八倍体小偃麦,表现出优异的黄矮病和锈病抗性[63-65]。中国科学院西北植物所利用长穗偃麦草与普通小麦杂交,育成了“小偃”系列八倍体[66]。山东农业大学利用普通小麦分别与长穗偃麦草、中间偃麦草杂交创制出一批八倍体小偃麦[67-68]。以上材料为偃麦草属优异基因向小麦转移起到了桥梁作用。
然而外源染色体上的优异基因通常携带连锁累赘。因此,遗传稳定且遗传累赘少的小偃麦异附加系、代换系、易位系和渐渗系,在育种实践中更容易利用。例如,东北师范大学利用“中”字系列八倍体小偃麦创制出了两套小冰麦异附加系,其中TAI-23高抗秆锈病,TAI-25高抗叶锈病[69]。异附加系WTA55苗期和成株期高抗条锈病,其抗性可能来源于长穗偃麦草第6部分同源群染色体[4]。小偃麦附加系小偃85高抗秆锈病Ug99,其抗性可能来源于长穗偃麦草第1部分同源群染色体[4, 70]。小偃麦5Ag(5D)代换系PY5814成株期高抗叶锈病和白粉病,连锁分析判断其抗性来源于5Ag染色体[37]。小偃麦易位系RWG33和RWG34携带抗秆锈病基因Sr43[43],易位系1-40和9-80携带抗叶锈病基因Lr19[30],易位系WTT26苗期近免疫叶锈病,其抗性基因来源于中间偃麦草7Js染色体[71]。小偃麦渐渗系SN0293-2和SN0293-7成株期高抗白粉病,其抗性可能来源于长穗偃麦草[72]。
偃麦草属植物具有很强的抗病性,早在20世纪50年代,以李振声院士为代表的国内科学家开始进行小麦与长穗偃麦草、中间偃麦草的远缘杂交研究。截至2025年底,已培育出多个抗病性状优良的小麦品种,为保护国家粮食安全作出突出贡献。其中,孙善澄[63]利用普通小麦与中间偃麦草一次杂交培育出龙麦1号、龙麦2号、龙麦3号等4个小麦新品种,用小偃麦中间类型与小麦回交育成高抗叶锈病的新品种新曙光8号和新曙光9号。李振声团队利用普通小麦与长穗偃麦草杂交培育出小偃6号,该品种以其高产、抗病、优质等特点,于1981年通过陕西省农作物品种审定委员会审定[73]。以小偃6号为直接或间接亲本育成的小麦品种已达80余个,累计推广面积超0.2亿hm2,增产小麦达750万t[73]。通过染色体工程和常规育种技术育成的小冰麦33,是高产、多抗、优质的春小麦品种,于1995年审定推广[8]。高优503具有高产、优质、综合抗性好等优点,分别于1997年、1998年通过陕西省和河北省品种审定委员会审定[9]。西农511分别于2015年和2018年通过陕西省和国家农作物品种审定委员会审定,中抗赤霉病,已定位2个可能来源于长穗偃麦草的抗赤霉病QTLs QFhb.nwafu-5B和QFhb.nwafu-7A[60]。2021年通过山东省审定的山农48,是首个携带抗赤霉病基因Fhb7的小麦新品种,该品种表现为中抗赤霉病。中科166携带二倍体长穗偃麦草Fhb7-7EL位点,中抗赤霉病,于2022年通过国家品种审定委员会审定[58]。在国外,澳大利亚育种家利用携带抗秆锈病基因Sr26的易位系9A-1育成了抗秆锈病新品种Eagle[74]。
加强偃麦草属植物抗病基因研究,对提高偃麦草属利用效率,推动小麦抗病育种具有重要意义。然而,偃麦草属植物基因组庞大,已挖掘的抗病基因只是冰山一角,偃麦草属抗病基因的研究和应用仍面临一些问题。一方面,偃麦草属植物中抗病基因挖掘不足、抗病基因的作用机制有待解析。例如,十倍体长穗偃麦草具有突出的条锈病抗性,但目前仅鉴定出1个抗病基因Yr69。虽然通过远缘杂交和分子染色体工程,已创制出多个高抗条锈病的小偃麦新种质,但其抗性来源有待进一步验证。在已鉴定出的抗病基因中,仅有3个抗秆锈病基因和1个抗赤霉病基因被克隆,且关于其作用机制研究甚少。另一方面,携带抗病基因的偃麦草染色体片段存在连锁累赘、远缘杂交后代优异性状不易稳定,导致偃麦草属抗病基因利用效率较低。目前,培育出携带已知偃麦草抗病基因的品种很少。因此,如何快速克隆偃麦草中的抗病基因,并将其高效应用于小麦抗病育种仍是当前偃麦草属抗病基因挖掘和利用的重点问题。
偃麦草染色体与小麦染色体不易发生染色体交换,利用常规图位克隆方法克隆抗病基因非常困难。但随着二倍体长穗偃麦草、中间偃麦草基因组测序完成,小麦基因组进入泛基因组研究时代[75],加之分子生物学和生物信息学发展迅速,新兴技术不断产生和升级,如MutRenSeq[10]、MutIsoSeq[76]、STAM[77]等,为利用比较基因组学、创制感病突变体等方法快速克隆偃麦草属抗病基因带来了新的机遇。在抗病基因育种利用方面,可将传统染色体工程技术与新兴技术手段相结合。如使用新型染色体工具材料,创制无遗传累赘、农艺性状好的新种质[78];利用快速加代技术、单倍体育种等,实现远缘杂交后代的快速纯合稳定等[79-80]。随着研究的不断深入,偃麦草属植物将在小麦遗传改良中体现其更高的应用价值,为小麦新一轮增产提质作出新的重要贡献。

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doi: 10.13430/j.cnki.jpgr.20250919003
  • 接收时间:2025-09-19
  • 首发时间:2026-09-30
  • 出版时间:2026-07-16
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    1山东农业大学农学院,泰安 271018
    2小麦育种全国重点实验室,山东 泰安 271018
    3山东农业大学未来技术学院(齐鲁学堂),泰安 271018

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杨国堂,研究方向为小偃麦分子染色体工程,E-mail:
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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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