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Article(id=1302192563860038392, tenantId=1146029695717560320, journalId=1301850032934322245, issueId=1302192562882761358, articleNumber=null, orderNo=null, doi=10.3864/j.issn.0578-1752.2026.16.007, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1774108800000, receivedDateStr=2026-03-22, revisedDate=null, revisedDateStr=null, acceptedDate=1780675200000, acceptedDateStr=2026-06-06, onlineDate=1788396500792, onlineDateStr=2026-09-03, pubDate=1786809600000, pubDateStr=2026-08-16, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1788396500792, onlineIssueDateStr=2026-09-03, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1788396500792, creator=13701087609, updateTime=1788396500792, updator=13701087609, issue=Issue{id=1302192562882761358, tenantId=1146029695717560320, journalId=1301850032934322245, year='2026', volume='59', issue='16', pageStart='3465', pageEnd='3698', issueExtLink='null', onlineDate='null', pubDate='1786809600000', pubDateStr='2026-08-16', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1788396500558, creator='13701087609', updateTime=1788405251849, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1302229268860264480, tenantId=1146029695717560320, journalId=1301850032934322245, issueId=1302192562882761358, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1302229268860264481, tenantId=1146029695717560320, journalId=1301850032934322245, issueId=1302192562882761358, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=3556, endPage=3576, ext={EN=ArticleExt(id=1302192566225625850, articleId=1302192563860038392, tenantId=1146029695717560320, journalId=1301850032934322245, language=EN, title=Advances in Mechanisms and Innovative Improvement Strategies for Wheat Disease Resistance, columnId=1302192564501766905, journalTitle=Scientia Agricultura Sinica, columnName=PLANT PROTECTION, runingTitle=null, highlight=null, articleAbstract=

Wheat (Triticum aestivum) remains vital to global food security, yet diverse pathogens constantly threaten its stable production. To address these threats, the identification and utilization of resistant genetic resources is the most effective and eco-friendly approach to manage disease epidemics. Based on a systematic review of the molecular mechanisms of wheat immunity, this paper compares the infection strategies of biotrophic and necrotrophic pathogens and elucidates the evolutionary arms race between wheat and its pathogens. Particular emphasis is placed on key immune regulatory mechanisms, including resistosome assembly and allele-specific evolution, modular cooperation mediated by kinase-integrated immune receptors, and host physiological homeostasis reprogramming driven by non-canonical resistance genes. In addition, the molecular basis by which pathogen effectors promote susceptibility through hijacking host immune pathways or exploiting hypersensitive response-associated cell death is discussed. Building upon these mechanistic insights, we summarize current strategies for resistance resource discovery and precision improvement, including the construction of panoramic resistance-gene atlases, the exploitation of novel resistance resources from wild relatives, and receptor optimization through targeted editing of key genetic loci. In response to the continuing evolution of pathogen populations and the increasing prevalence of multiple concurrent diseases, we further propose several emerging directions for resistance improvement, including the decoupling of immune activation from cell death, the evolution-guided design of universal immune receptors, and the establishment of multi-kingdom immune ecological barriers. These concepts provide a theoretical framework and technical foundation for the rational design of broad-spectrum and durable disease resistance in wheat.

, authors=JianHui WU1, 3, QingDong ZENG2, 3, ShengJie LIU2, 3, XiaoJie WANG2, 3, DeJun HAN1, 3, ZhenSheng KANG2, 3, authorsList=JianHui WU, QingDong ZENG, ShengJie LIU, XiaoJie WANG, DeJun HAN, ZhenSheng KANG, authorCompany=null, 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=1302192567760741124, articleId=1302192563860038392, tenantId=1146029695717560320, journalId=1301850032934322245, language=CN, title=小麦抗病机制研究进展与抗性创新利用策略, columnId=1302192566309511931, journalTitle=中国农业科学, columnName=植物保护, runingTitle=null, highlight=null, articleAbstract=

小麦（Triticum aestivum）对全球粮食安全至关重要，但其生产持续受到各类病原菌的侵害。发掘利用抗病资源是应对病害威胁、实现绿色防控的首选途径。本文在系统梳理小麦抗病分子机制研究进展的基础上，通过比较活体营养与死体营养型病原菌侵染特征，阐明了小麦与病原菌协同进化下的攻防演替策略；重点分析了抗病小体组装与等位基因特异性演变、激酶融合蛋白介导的模块化协作、非经典基因驱动的寄主生理稳态重构等免疫调控机制，并阐释了效应子通过劫持寄主免疫通路或利用超敏反应诱导感病的分子基础。基于上述免疫机制认识，总结了当前抗病资源发掘与精准改良的重要策略，包括构建抗病基因全景导航图、深度发掘野生近缘种资源以及通过关键遗传位点编辑实现免疫受体定向优化等。面对未来病原菌持续演化与多病害复合发生趋势，提出以防御激活与细胞死亡解耦、进化导向的通用免疫受体设计及多界免疫生态屏障构建为代表的新型抗性改良方向，为实现小麦广谱、持久抗性的理性设计提供理论依据与技术参考。

, authors=吴建辉1, 3, 曾庆东2, 3, 刘胜杰2, 3, 王晓杰2, 3, 韩德俊1, 3, 康振生2, 3, authorsList=吴建辉, 曾庆东, 刘胜杰, 王晓杰, 韩德俊, 康振生, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=
吴建辉，E-mail：
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Dissection of genetic structure and genetic basis of important traits for the new wheat lines based on 0.1K prospect and 16K background arrays[J]. Journal of Triticeae Crops, 2025, 45(1): 52-62. (in Chinese), articleTitle=Dissection of genetic structure and genetic basis of important traits for the new wheat lines based on 0.1K prospect and 16K background arrays, refAbstract=null), Reference(id=1302192590305125318, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, doi=null, pmid=null, pmcid=null, year=2023, volume=4, issue=3, pageStart=100549, pageEnd=null, url=null, language=null, rfNumber=[114], rfOrder=118, authorNames=Jiao C, Hao C, Li T, Bohra A, Wang L, Hou J, Liu H, Liu H, Zhao J, Wang Y, Liu Y, Wang Z, Jing X, Wang X, Varshney R K, Fu J, Zhang X, journalName=Plant Communications, refType=null, unstructuredReference=Jiao C, Hao C, Li T, Bohra A, Wang L, Hou J, Liu H, Liu H, Zhao J, Wang Y, Liu Y, Wang Z, Jing X, Wang X, Varshney R K, Fu J, Zhang X. Fast integration and accumulation of beneficial breeding alleles through an AB-NAMIC strategy in wheat[J]. Plant Communications, 2023, 4(3): 100549., articleTitle=Fast integration and accumulation of beneficial breeding alleles through an AB-NAMIC strategy in wheat, refAbstract=null)], funds=[Fund(id=1302192573125256005, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, awardId=32272088, language=CN, fundingSource=国家自然科学基金(32272088), fundOrder=null, country=null), Fund(id=1302192573213336390, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, awardId=32225041, language=CN, fundingSource=国家自然科学基金(32225041), fundOrder=null, country=null), Fund(id=1302192573293028167, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, awardId=32472103, language=CN, fundingSource=国家自然科学基金(32472103), fundOrder=null, country=null), Fund(id=1302192573381108552, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, awardId=32561143296, language=CN, fundingSource=国家自然科学基金(32561143296), fundOrder=null, country=null), Fund(id=1302192573460800329, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, awardId=2021YFD1401000, language=CN, fundingSource=国家重点研发计划(2021YFD1401000), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1302192568054342405, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, xref=1, ext=[AuthorCompanyExt(id=1302192568066925318, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568054342405, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1 College of Agronomy, Northwest A&F University, Yangling 712100, Shaanxi), AuthorCompanyExt(id=1302192568079508231, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568054342405, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1 西北农林科技大学农学院， 陕西杨凌 712100)]), AuthorCompany(id=1302192568255669000, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, xref=2, ext=[AuthorCompanyExt(id=1302192568264057609, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568255669000, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2 College of Plant Protection, Northwest A&F University, Yangling 712100, Shaanxi), AuthorCompanyExt(id=1302192568272446218, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568255669000, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2 西北农林科技大学植物保护学院， 陕西杨凌 712100)]), AuthorCompany(id=1302192568347943691, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, xref=3, ext=[AuthorCompanyExt(id=1302192568356332300, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568347943691, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3 State Key Laboratory of Crop Stress Resistance and High-Efficiency Production, Yangling 712100, Shaanxi), AuthorCompanyExt(id=1302192568364720909, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, companyId=1302192568347943691, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3 作物抗逆与高效生产全国重点实验室， 陕西杨凌 712100)])], figs=[ArticleFig(id=1302192572085068603, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=EN, label=Fig. 1, caption=Disease symptoms of major wheat diseases

Stripe rust, leaf rust, stem rust and powdery mildew are biotrophic diseases, mainly occurring on leaves and stems. The rest are necrotrophic diseases. Among them, fusarium head blight and wheat blast occur on spikes, septoria tritici blotch infects leaves, and sharp eyespot as well as fusarium crown rot prevail on stems. The pictures of wheat blast and septoria tritici blotch are referenced from Singh et al[15-16]

, figureFileSmall=qyx6OAWzpQGv+uCUQHDuZQ==, figureFileBig=/N3WJ+qsNgZW/yV7ururfQ==, tableContent=null), ArticleFig(id=1302192572164760380, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=CN, label=图1, caption=小麦主要病害发病特征

条锈病、叶锈病、秆锈病、白粉病为活体营养型病害，主要发生在叶部和茎部；其他为死体营养型病害，其中赤霉病和麦瘟病发生在穗部，叶枯病发生在叶部，纹枯病和茎基腐病发生在茎部。麦瘟病和叶枯病图片参考Singh等[15-16]

, figureFileSmall=qyx6OAWzpQGv+uCUQHDuZQ==, figureFileBig=/N3WJ+qsNgZW/yV7ururfQ==, tableContent=null), ArticleFig(id=1302192572349309757, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=EN, label=Fig. 2, caption=Types of wheat disease resistance, figureFileSmall=7jUKS3jtxpH++1+G+kxs1A==, figureFileBig=BJ8TYVeBrBTCXODbbtkLMg==, tableContent=null), ArticleFig(id=1302192572420612926, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=CN, label=图2, caption=小麦抗病性类型, figureFileSmall=7jUKS3jtxpH++1+G+kxs1A==, figureFileBig=BJ8TYVeBrBTCXODbbtkLMg==, tableContent=null), ArticleFig(id=1302192572487721791, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=EN, label=Fig. 3, caption=Structural diversity and mediated immune defense mechanisms of wheat disease resistance genes, figureFileSmall=QuSXlgB8Ioucoxg9fukRtg==, figureFileBig=zuBllVm72ebvdy7XBmEipQ==, tableContent=null), ArticleFig(id=1302192572559024960, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=CN, label=图3, caption=小麦抗病基因的结构多样性及其介导的免疫防御机制, figureFileSmall=QuSXlgB8Ioucoxg9fukRtg==, figureFileBig=zuBllVm72ebvdy7XBmEipQ==, tableContent=null), ArticleFig(id=1302192572621939521, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=EN, label=Fig. 4, caption=Future strategies for wheat disease resistance breeding, figureFileSmall=9lM/6gJDYd6taSJo6EhZrA==, figureFileBig=qdOLGLBmQ63AmPOHnOxX5w==, tableContent=null), ArticleFig(id=1302192572718408514, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=CN, label=图4, caption=未来小麦抗病育种策略, figureFileSmall=9lM/6gJDYd6taSJo6EhZrA==, figureFileBig=qdOLGLBmQ63AmPOHnOxX5w==, tableContent=null), ArticleFig(id=1302192572814877507, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=EN, label=Table 1, caption=

Cloned disease resistance genes in wheat

, figureFileSmall=null, figureFileBig=null, tableContent=
基因Gene 蛋白Protein 位点Loci 供体Donor 病害Disease
Yr5a/b/x BED-NLR 2BL 斯贝尔脱小麦T. spelta；
普通小麦T. aestivum		 条锈病Stripe rust (Pst)
Yr61 CNL pair 7BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr7 BED-NLR 2BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr9 CNL 1RS 黑麦S. cereale 条锈病Stripe rust (Pst)
YrNAM (Yr10) NAM-ZnF-BED 1BS 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr15 (WTK1) KD-PKD 1BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
Yr26 DTMP 1BL 硬粒小麦T. durum 条锈病Stripe rust (Pst)
Yr272 CNL 2BS 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr28/AS2388R; YrAet672 CNL 4DS 山羊草Ae. tauschii 条锈病Stripe rust (Pst)
Yr36 (WKS1) KD-START 6BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
Yr84/TD121 CNL and NL pair 1BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
YrKB PH-START-EDR2 7BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
YrU1 ANK-NLR-WRKY 5AL 乌拉尔图小麦T. urartu 条锈病Stripe rust (Pst)
Lr1 CNL 5DL 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr10-Rga2 CNL and CNL pair 1AS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr13 (Ne2)2 CNL 2BS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr14a Ankyrin TM protein 7BL 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr21 CNL 1DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr22a CNL 2DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr30 CNL 4AS 硬粒小麦T. durum 叶锈病Leaf rust (Pt)
Lr39 KD-KD-MSP-WD40 2DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr42 CNL 1DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr47 CNL 7S#1S 斯贝尔脱山羊草Ae. speltoides 叶锈病Leaf rust (Pt)
Lr9/Lr58 (WTK6)3 KD-PKD-vWA 6BL 小伞山羊草Ae. umbellulata 叶锈病Leaf rust (Pt)
Rga2 NLR-ID 1DS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
AcRLK2P-1 LRR-RK 2PL 冰草Ag. cristatum 叶锈病Leaf rust (Pt)
Sr6 BED-NLR 2DS 普通小麦T. aestivum 秆锈病Stem rust (Pgt)
Sr9 CNL 2BL 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Sr13 CNL 6AL 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Sr21 CNL 2AmL 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr22a/b4 CNL 7AbL; 7AmL 野生一粒小麦T. boeoticum；
一粒小麦T. monococcum		 秆锈病Stem rust (Pgt)
Sr26 CNL 6Ae#1L 十倍体长穗偃麦草Th. ponticum 秆锈病Stem rust (Pgt)
Sr27 CNL 3RS 黑麦S. cereale 秆锈病Stem rust (Pgt)
Sr33 CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr35 CNL 3AmL 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr43 KD-DUFs 7EL 二倍体长穗偃麦草Th. elongatum 秆锈病Stem rust (Pgt)
Sr45 CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr46 CNL 2DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr50 CNL 1RS 黑麦S. cereale 秆锈病Stem rust (Pgt)
Sr60 (WTK2) KD-KD 5AmS 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr61 CNL 6Ae#3L 十倍体长穗偃麦草Th. ponticum 秆锈病Stem rust (Pgt)
Sr62 (WTK5) KD-PKD 1SshS 沙融山羊草Ae. sharonensis 秆锈病Stem rust (Pgt)
Sr66 (TA1662) CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr8155B1 CNL 6AS 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Pm1a CNL 7AL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm2 CNL 5DS 山羊草Ae. tauschii 白粉病Powdery mildew (Bgt)
Pm3a/b/d CNL 1AS 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm4b5 KD-MCTP 2AL 波斯小麦T. carthlicum 白粉病Powdery mildew (Bgt)
Pm5e1 CNL pair 7BL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm6/52 CNL 2GL 提莫菲维小麦T. timopheevii 白粉病Powdery mildew (Bgt)
Pm8, Pm17 CNL 1RS 黑麦S. cereale 白粉病Powdery mildew (Bgt)
Pm12 CNL 6SS 斯贝尔脱山羊草Ae. speltoides 白粉病Powdery mildew (Bgt)
Pm13 HeLo-KD 3SlS 高大山羊草Ae. longissima 白粉病Powdery mildew (Bgt)
Pm21 CNL 6VS 簇毛麦D. villosum 白粉病Powdery mildew (Bgt)
Pm24 (WTK3)6 KD-PKD 1DS 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm26 CNL pair 3BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm36 (WTK7) KD-KD-TM 5BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm374 CNL 7AL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm41 CNL 3BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
SuPm55-Pm55a/b CNL 5VS 簇毛麦D. villosum 白粉病Powdery mildew (Bgt)
Pm57 (WTK6b)3 KD-PKD-vWA 2SL 西尔斯山羊草Ae. searsii 白粉病Powdery mildew (Bgt)
Pm60a/b; MlIW18/172 CNL 7AL 乌拉尔图小麦T. urartu；
野生二粒小麦T. dicoccoides		 白粉病Powdery mildew (Bgt)
Pm68/WR183/MlIW39 CNL pair 6BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm69 CNL 6BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm6Sl BED-NLR 6SlL 高大山羊草Ae. longissima 白粉病Powdery mildew (Bgt)
PmAeu1 CNL 2UL 小伞山羊草Ae. umbellulata 白粉病Powdery mildew (Bgt)
PmTR1, PmTR3 CNL 6RS 黑麦S. cereale 白粉病Powdery mildew (Bgt)
WTK4 KD-PKD 7DS 山羊草Ae. tauschii 白粉病Powdery mildew (Bgt)
Yr18/Lr34/Sr57/Pm38 ABC transporter 7DS 普通小麦T. aestivum 兼抗Multiple diseases resistance (Pt/Pst/Pgt/Bgt)
Yr46/Lr67/Sr55/Pm46 Hexose-proton symporter 4DL 普通小麦T. aestivum 兼抗Multiple diseases resistance (Pt/Pst/Pgt/Bgt)
Lr85/Yr87 CNL 6SshS/6SlS 沙融山羊草Ae. sharonensis；
高大山羊草Ae. longissima		 兼抗Multiple diseases resistance (Pt/Pst)
Fhb1 His 3BS 普通小麦T. aestivum 赤霉病Fusarium head blight (Fg)
Fhb7 GST 7EL 二倍体长穗偃麦草Th. elongatum 赤霉病Fusarium head blight (Fg)
Stb15 LecRK 6AS 普通小麦T. aestivum 叶枯病Septoria tritici blotch (Zt)
Stb16q CRK 3DS 山羊草Ae. tauschii 叶枯病Septoria tritici blotch (Zt)
Stb6 WAK 3AS 普通小麦T. aestivum 叶枯病Septoria tritici blotch (Zt)
Snn1 WAK 1BS 普通小麦T. aestivum 颖枯病Septoria nodorum blotch (Pn)
Snn3 KD-MSP 5DS 山羊草Ae. tauschii 颖枯病Septoria nodorum blotch (Pn)
Tsn1 KD-NBS-LRR 5BL 普通小麦T. aestivum 褐斑病Tan spot (Bs)
Rmg75 KD-MCTP 2AL 栽培二粒小麦T. dicoccum 麦瘟病Wheat blast (Po)
Rmg8 KD-MCTP 2BL 普通小麦T. aestivum 麦瘟病Wheat blast (Po)
Rwt3 CNL 1DS 普通小麦T. aestivum 麦瘟病Wheat blast (Po)
RWT46 KD-PKD 1DS 山羊草Ae. tauschii 麦瘟病Wheat blast (Po)
), ArticleFig(id=1302192572965872452, tenantId=1146029695717560320, journalId=1301850032934322245, articleId=1302192563860038392, language=CN, label=表1, caption=

已克隆的小麦抗病基因

, figureFileSmall=null, figureFileBig=null, tableContent=
基因Gene 蛋白Protein 位点Loci 供体Donor 病害Disease
Yr5a/b/x BED-NLR 2BL 斯贝尔脱小麦T. spelta；
普通小麦T. aestivum		 条锈病Stripe rust (Pst)
Yr61 CNL pair 7BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr7 BED-NLR 2BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr9 CNL 1RS 黑麦S. cereale 条锈病Stripe rust (Pst)
YrNAM (Yr10) NAM-ZnF-BED 1BS 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr15 (WTK1) KD-PKD 1BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
Yr26 DTMP 1BL 硬粒小麦T. durum 条锈病Stripe rust (Pst)
Yr272 CNL 2BS 普通小麦T. aestivum 条锈病Stripe rust (Pst)
Yr28/AS2388R; YrAet672 CNL 4DS 山羊草Ae. tauschii 条锈病Stripe rust (Pst)
Yr36 (WKS1) KD-START 6BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
Yr84/TD121 CNL and NL pair 1BS 野生二粒小麦T. dicoccoides 条锈病Stripe rust (Pst)
YrKB PH-START-EDR2 7BL 普通小麦T. aestivum 条锈病Stripe rust (Pst)
YrU1 ANK-NLR-WRKY 5AL 乌拉尔图小麦T. urartu 条锈病Stripe rust (Pst)
Lr1 CNL 5DL 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr10-Rga2 CNL and CNL pair 1AS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr13 (Ne2)2 CNL 2BS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr14a Ankyrin TM protein 7BL 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
Lr21 CNL 1DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr22a CNL 2DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr30 CNL 4AS 硬粒小麦T. durum 叶锈病Leaf rust (Pt)
Lr39 KD-KD-MSP-WD40 2DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr42 CNL 1DS 山羊草Ae. tauschii 叶锈病Leaf rust (Pt)
Lr47 CNL 7S#1S 斯贝尔脱山羊草Ae. speltoides 叶锈病Leaf rust (Pt)
Lr9/Lr58 (WTK6)3 KD-PKD-vWA 6BL 小伞山羊草Ae. umbellulata 叶锈病Leaf rust (Pt)
Rga2 NLR-ID 1DS 普通小麦T. aestivum 叶锈病Leaf rust (Pt)
AcRLK2P-1 LRR-RK 2PL 冰草Ag. cristatum 叶锈病Leaf rust (Pt)
Sr6 BED-NLR 2DS 普通小麦T. aestivum 秆锈病Stem rust (Pgt)
Sr9 CNL 2BL 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Sr13 CNL 6AL 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Sr21 CNL 2AmL 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr22a/b4 CNL 7AbL; 7AmL 野生一粒小麦T. boeoticum；
一粒小麦T. monococcum		 秆锈病Stem rust (Pgt)
Sr26 CNL 6Ae#1L 十倍体长穗偃麦草Th. ponticum 秆锈病Stem rust (Pgt)
Sr27 CNL 3RS 黑麦S. cereale 秆锈病Stem rust (Pgt)
Sr33 CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr35 CNL 3AmL 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr43 KD-DUFs 7EL 二倍体长穗偃麦草Th. elongatum 秆锈病Stem rust (Pgt)
Sr45 CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr46 CNL 2DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr50 CNL 1RS 黑麦S. cereale 秆锈病Stem rust (Pgt)
Sr60 (WTK2) KD-KD 5AmS 一粒小麦T. monococcum 秆锈病Stem rust (Pgt)
Sr61 CNL 6Ae#3L 十倍体长穗偃麦草Th. ponticum 秆锈病Stem rust (Pgt)
Sr62 (WTK5) KD-PKD 1SshS 沙融山羊草Ae. sharonensis 秆锈病Stem rust (Pgt)
Sr66 (TA1662) CNL 1DS 山羊草Ae. tauschii 秆锈病Stem rust (Pgt)
Sr8155B1 CNL 6AS 硬粒小麦T. durum 秆锈病Stem rust (Pgt)
Pm1a CNL 7AL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm2 CNL 5DS 山羊草Ae. tauschii 白粉病Powdery mildew (Bgt)
Pm3a/b/d CNL 1AS 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm4b5 KD-MCTP 2AL 波斯小麦T. carthlicum 白粉病Powdery mildew (Bgt)
Pm5e1 CNL pair 7BL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm6/52 CNL 2GL 提莫菲维小麦T. timopheevii 白粉病Powdery mildew (Bgt)
Pm8, Pm17 CNL 1RS 黑麦S. cereale 白粉病Powdery mildew (Bgt)
Pm12 CNL 6SS 斯贝尔脱山羊草Ae. speltoides 白粉病Powdery mildew (Bgt)
Pm13 HeLo-KD 3SlS 高大山羊草Ae. longissima 白粉病Powdery mildew (Bgt)
Pm21 CNL 6VS 簇毛麦D. villosum 白粉病Powdery mildew (Bgt)
Pm24 (WTK3)6 KD-PKD 1DS 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm26 CNL pair 3BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm36 (WTK7) KD-KD-TM 5BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm374 CNL 7AL 普通小麦T. aestivum 白粉病Powdery mildew (Bgt)
Pm41 CNL 3BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
SuPm55-Pm55a/b CNL 5VS 簇毛麦D. villosum 白粉病Powdery mildew (Bgt)
Pm57 (WTK6b)3 KD-PKD-vWA 2SL 西尔斯山羊草Ae. searsii 白粉病Powdery mildew (Bgt)
Pm60a/b; MlIW18/172 CNL 7AL 乌拉尔图小麦T. urartu；
野生二粒小麦T. dicoccoides		 白粉病Powdery mildew (Bgt)
Pm68/WR183/MlIW39 CNL pair 6BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm69 CNL 6BL 野生二粒小麦T. dicoccoides 白粉病Powdery mildew (Bgt)
Pm6Sl BED-NLR 6SlL 高大山羊草Ae. longissima 白粉病Powdery mildew (Bgt)
PmAeu1 CNL 2UL 小伞山羊草Ae. umbellulata 白粉病Powdery mildew (Bgt)
PmTR1, PmTR3 CNL 6RS 黑麦S. cereale 白粉病Powdery mildew (Bgt)
WTK4 KD-PKD 7DS 山羊草Ae. tauschii 白粉病Powdery mildew (Bgt)
Yr18/Lr34/Sr57/Pm38 ABC transporter 7DS 普通小麦T. aestivum 兼抗Multiple diseases resistance (Pt/Pst/Pgt/Bgt)
Yr46/Lr67/Sr55/Pm46 Hexose-proton symporter 4DL 普通小麦T. aestivum 兼抗Multiple diseases resistance (Pt/Pst/Pgt/Bgt)
Lr85/Yr87 CNL 6SshS/6SlS 沙融山羊草Ae. sharonensis；
高大山羊草Ae. longissima		 兼抗Multiple diseases resistance (Pt/Pst)
Fhb1 His 3BS 普通小麦T. aestivum 赤霉病Fusarium head blight (Fg)
Fhb7 GST 7EL 二倍体长穗偃麦草Th. elongatum 赤霉病Fusarium head blight (Fg)
Stb15 LecRK 6AS 普通小麦T. aestivum 叶枯病Septoria tritici blotch (Zt)
Stb16q CRK 3DS 山羊草Ae. tauschii 叶枯病Septoria tritici blotch (Zt)
Stb6 WAK 3AS 普通小麦T. aestivum 叶枯病Septoria tritici blotch (Zt)
Snn1 WAK 1BS 普通小麦T. aestivum 颖枯病Septoria nodorum blotch (Pn)
Snn3 KD-MSP 5DS 山羊草Ae. tauschii 颖枯病Septoria nodorum blotch (Pn)
Tsn1 KD-NBS-LRR 5BL 普通小麦T. aestivum 褐斑病Tan spot (Bs)
Rmg75 KD-MCTP 2AL 栽培二粒小麦T. dicoccum 麦瘟病Wheat blast (Po)
Rmg8 KD-MCTP 2BL 普通小麦T. aestivum 麦瘟病Wheat blast (Po)
Rwt3 CNL 1DS 普通小麦T. aestivum 麦瘟病Wheat blast (Po)
RWT46 KD-PKD 1DS 山羊草Ae. tauschii 麦瘟病Wheat blast (Po)
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小麦抗病机制研究进展与抗性创新利用策略
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吴建辉 1, 3 , 曾庆东 2, 3 , 刘胜杰 2, 3 , 王晓杰 2, 3 , 韩德俊 1, 3 , 康振生 2, 3
中国农业科学 | 植物保护 2026,59(16): 3556-3576
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中国农业科学 |植物保护 2026 , 59 (16) : 3556 -3576
小麦抗病机制研究进展与抗性创新利用策略
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吴建辉1, 3 , 曾庆东2, 3, 刘胜杰2, 3, 王晓杰2, 3, 韩德俊1, 3, 康振生2, 3
作者信息
  • 1 西北农林科技大学农学院， 陕西杨凌 712100
  • 2 西北农林科技大学植物保护学院， 陕西杨凌 712100
  • 3 作物抗逆与高效生产全国重点实验室， 陕西杨凌 712100
通讯作者:
吴建辉，E-mail：
Advances in Mechanisms and Innovative Improvement Strategies for Wheat Disease Resistance
JianHui WU1, 3 , QingDong ZENG2, 3, ShengJie LIU2, 3, XiaoJie WANG2, 3, DeJun HAN1, 3, ZhenSheng KANG2, 3
Affiliations
  • 1 College of Agronomy, Northwest A&F University, Yangling 712100, Shaanxi
  • 2 College of Plant Protection, Northwest A&F University, Yangling 712100, Shaanxi
  • 3 State Key Laboratory of Crop Stress Resistance and High-Efficiency Production, Yangling 712100, Shaanxi
出版时间: 2026-08-16 doi: 10.3864/j.issn.0578-1752.2026.16.007
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小麦（Triticum aestivum）对全球粮食安全至关重要，但其生产持续受到各类病原菌的侵害。发掘利用抗病资源是应对病害威胁、实现绿色防控的首选途径。本文在系统梳理小麦抗病分子机制研究进展的基础上，通过比较活体营养与死体营养型病原菌侵染特征，阐明了小麦与病原菌协同进化下的攻防演替策略；重点分析了抗病小体组装与等位基因特异性演变、激酶融合蛋白介导的模块化协作、非经典基因驱动的寄主生理稳态重构等免疫调控机制，并阐释了效应子通过劫持寄主免疫通路或利用超敏反应诱导感病的分子基础。基于上述免疫机制认识，总结了当前抗病资源发掘与精准改良的重要策略，包括构建抗病基因全景导航图、深度发掘野生近缘种资源以及通过关键遗传位点编辑实现免疫受体定向优化等。面对未来病原菌持续演化与多病害复合发生趋势，提出以防御激活与细胞死亡解耦、进化导向的通用免疫受体设计及多界免疫生态屏障构建为代表的新型抗性改良方向，为实现小麦广谱、持久抗性的理性设计提供理论依据与技术参考。

小麦  /  抗病基因  /  抗/感病机制  /  抗病性改良

Wheat (Triticum aestivum) remains vital to global food security, yet diverse pathogens constantly threaten its stable production. To address these threats, the identification and utilization of resistant genetic resources is the most effective and eco-friendly approach to manage disease epidemics. Based on a systematic review of the molecular mechanisms of wheat immunity, this paper compares the infection strategies of biotrophic and necrotrophic pathogens and elucidates the evolutionary arms race between wheat and its pathogens. Particular emphasis is placed on key immune regulatory mechanisms, including resistosome assembly and allele-specific evolution, modular cooperation mediated by kinase-integrated immune receptors, and host physiological homeostasis reprogramming driven by non-canonical resistance genes. In addition, the molecular basis by which pathogen effectors promote susceptibility through hijacking host immune pathways or exploiting hypersensitive response-associated cell death is discussed. Building upon these mechanistic insights, we summarize current strategies for resistance resource discovery and precision improvement, including the construction of panoramic resistance-gene atlases, the exploitation of novel resistance resources from wild relatives, and receptor optimization through targeted editing of key genetic loci. In response to the continuing evolution of pathogen populations and the increasing prevalence of multiple concurrent diseases, we further propose several emerging directions for resistance improvement, including the decoupling of immune activation from cell death, the evolution-guided design of universal immune receptors, and the establishment of multi-kingdom immune ecological barriers. These concepts provide a theoretical framework and technical foundation for the rational design of broad-spectrum and durable disease resistance in wheat.

wheat  /  disease resistance gene  /  resistance and susceptibility mechanism  /  improvement of disease resistance
吴建辉, 曾庆东, 刘胜杰, 王晓杰, 韩德俊, 康振生. 小麦抗病机制研究进展与抗性创新利用策略. 中国农业科学, 2026 , 59 (16) : 3556 -3576 . DOI: 10.3864/j.issn.0578-1752.2026.16.007
JianHui WU, QingDong ZENG, ShengJie LIU, XiaoJie WANG, DeJun HAN, ZhenSheng KANG. Advances in Mechanisms and Innovative Improvement Strategies for Wheat Disease Resistance[J]. Scientia Agricultura Sinica, 2026 , 59 (16) : 3556 -3576 . DOI: 10.3864/j.issn.0578-1752.2026.16.007
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0 引言
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小麦（Triticum aestivum）是全球最重要的粮食作物之一，在保障粮食安全和人类营养供给中发挥着重要作用。自1961年以来，全球小麦收获面积始终稳定在约2.2亿公顷，过去10年间其年均产量占全球谷物总产量的28.2%，凸显了其在世界粮食生产体系中的重要地位[1]。作为异源六倍体作物，小麦经历两次杂交和多倍化事件形成了庞大而复杂的基因组（AABBDD，约16 Gb），其高度重复的序列结构为抗性基因的发掘、功能解析及遗传改良带来了巨大挑战。
据预测，到2050年全球人口将达到91亿，为满足不断增长的粮食需求，全球粮食产量需在现有基础上提高约70%[2]。然而，病虫害仍是影响小麦稳产高产的最主要限制因素。据统计，全球小麦因病虫害导致的年均产量损失约为21.5%，其中病害造成的损失占比高达19.8%[3]。因此，在保障粮食安全和应对病害威胁的双重需求驱动下，如何培育兼具广谱性和持久性的抗病品种，已成为现代小麦育种面临的重要科学问题。
1 病原菌快速演化下的小麦抗性改良挑战
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小麦稳产面临的核心挑战在于病原菌毒力演化与寄主抗性改良之间长期存在的速率失衡。一方面，病原菌群体具有较强的遗传变异能力和环境适应能力，在全球气候变化与农业生态系统持续演变的背景下，其演化速度不断加快，传播范围持续扩大。例如，小麦条锈病菌等病原菌表现出更强的高温适应性，导致其地理分布与危害区域持续扩张[4-9]；同时，病原菌毒性小种的快速更迭常在短时间内克服品种所携带的单一抗性基因，引发病害流行[10]；受耕作制度调整和生态环境变化的影响，赤霉病、茎基腐病等原本局部发生的病害逐渐演变为影响广泛的重要灾害[11-12]。另一方面，抗病基因从发现、克隆、功能解析到育种利用往往需要较长周期，使得抗性改良进程难以同步应对病原菌群体结构和毒性谱的快速变化[13]。这种病原菌毒力快速演化与寄主抗性改良相对滞后之间的动态失衡，已成为当前制约小麦持续稳产的重要瓶颈。这一现实挑战表明，仅依赖零散抗性位点的发掘已难以满足持久抗病育种的需求。尽管过去10余年间，随着基因组学、群体遗传学和分子生物学技术的快速发展，研究者已从小麦及其野生近缘种中发掘并克隆了大量抗病（resistance，R）基因[14]，但目前多数研究仍聚焦于单个位点的功能解析，对其在复杂免疫网络中的作用模式认识仍相对有限。因此，亟需从植物免疫机制层面深入解析抗病性的分子基础，厘清不同抗病因子的作用方式及其相互关系，将分散的抗性资源转化为可精准利用的遗传资源，进而构建起体系化的抗性屏障。这已成为突破现有认知局限、实现小麦持久抗病性改良的优选之路。
本综述围绕小麦抗病机制解析与抗性资源利用策略展开讨论。首先比较活体营养型和死体营养型病原菌的侵染特征及其病害发展规律，解析针对不同病原生活方式下的免疫调控逻辑。随后系统总结已克隆抗病位点的作用机制，重点讨论抗病小体组装与等位基因功能分化、激酶融合蛋白介导的协同免疫以及非经典抗病基因驱动的寄主稳态重构等关键机制，并阐释抗病蛋白在应对不同侵染策略时表现出的功能分化。进一步结合抗病基因全景导航图、野生近缘种资源发掘及免疫受体工程等研究进展，探讨抗性资源的高效利用与精准改良路径。最后，围绕多病害协同防控、抗性与产量协同提升以及植物微生物组辅助抗病等前沿方向，展望小麦抗性设计育种的发展趋势。
2 小麦主要病原菌的侵染特征与寄主防御类型
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2.1 小麦主要病原菌的营养类型与侵染策略
小麦在其全生命周期中面临着多种病原菌引发的病害威胁。根据营养利用方式和攻击策略的不同，这些病原菌可分为活体营养型和死体营养型，其发生重心随植株发育进程呈现出明显的时空推移。
生长前期及营养生长期是典型活体营养型病原菌的集中侵染期。此类病原菌以条锈菌（Puccinia striiformis f. sp. tritici，Pst）、叶锈菌（Puccinia triticina，Pt）、秆锈菌（Puccinia graminis f. sp. tritici，Pgt）及白粉菌（Blumeria graminis f. sp. tritici，Bgt）为代表，其生存高度依赖于活体寄主细胞（图1）。基因组学研究显示，此类病原菌普遍丢失了编码细胞壁降解酶和次级代谢物的基因，以最大限度地减少对寄主细胞壁的破坏，从而避免引发寄主的强烈免疫响应。取而代之的是，它们通过特化的吸器向寄主内部输送大量效应蛋白，精密地抑制防御机制并掠夺养分。这种通过减少免疫激活相关分子并强化效应子分泌能力形成的协同适应策略，构成了其高度特化的侵染方式，使其能在维持寄主细胞活性的前提下实现长期寄生[17]。这些病害主要危害叶片和茎秆，其夏孢子或分生孢子可借助气流远距离传播，并在小麦生长季早期快速扩散、诱发大规模流行。
随着植株进入花期至灌浆期，由死体营养型病原菌引起的侵染压力显著增加。其中以禾谷镰孢（Fusarium graminearum，Fg）引发的赤霉病、假禾谷镰孢（F. pseudograminearum，Fp）引发的茎基腐病、禾谷丝核菌（Rhizoctonia cerealis，Rc）引发的纹枯病以及主要由壳针孢菌（Zymoseptoria tritici，Zt）引起的叶枯病最为典型（图1）。这些病害往往在小麦生殖生长的关键期或生理衰老阶段暴发。与活体营养菌不同，死体营养菌主要依赖细胞死亡诱导和组织降解等机制完成侵染：其基因组中含有大量编码细胞壁降解酶、诱导细胞死亡因子和活性氧产生相关的基因，它们通过分泌脱氧雪腐镰刀菌烯醇（DON）、ToxA等毒素或降解酶，主动诱发寄主细胞发生大规模死亡，从而制造出利于其生存的坏死环境[18-20]。除造成严重的产量损失外，这些病原菌产生的霉菌毒素还对粮食和饲料安全构成持续性威胁[21]。
2.2 小麦抗性的分类特征
面对病原菌复杂多样的侵染策略，小麦逐步进化形成了由基础免疫与特异性免疫协同调控的抗病体系。该体系并非相互独立的防御层级，两类免疫机制之间存在广泛互作与交叉调控；其核心特征表现为不同小麦品种对病原菌不同小种或分离物呈现显著差异的防御响应，从而形成丰富的种质抗性变异。自Biffen[22]首次证实小麦对条锈病的抗性符合孟德尔遗传规律以来，抗性基因的挖掘与鉴定一直是小麦抗病育种的基石。通常情况下，寄主抗性可分为两类：一类是由主效R基因控制的质量抗性，表现为典型的全生育期抗性（all-stage resistance，ASR）或小种特异性免疫（race-specific resistance）（图2），此类抗性多符合Flor的“基因对基因”假说[23]，即寄主受体蛋白通过识别病原菌的无毒效应子诱发超敏反应（hypersensitive response，HR），对应于效应子触发的免疫（effector-triggered immunity，ETI）[24]。虽然此类抗性水平高，但因其遗传基础单一，容易随着病原菌毒性小种的演替而被突破。特别是在应对不同营养类型的病原体时，编码核苷酸结合位点与亮氨酸富集重复类型的受体蛋白（nucleotide binding domain leucine-rich-repeat-containing receptor，NLR）存在明显不同的作用模式：在抵抗活体营养型病害时作为核心免疫受体介导高效的防卫反应，而在面对死体营养型病害时，其介导的HR反应反而会被病原菌劫持以辅助侵染[20]。这种抗性机制的二元性，决定了小麦在应对复杂病害环境时必须依赖更多元化的免疫机制。
另一类是由多个微效基因或数量性状位点（QTL）控制的数量抗性，常表现为非小种特异性（nonrace- specific resistance）或成株期抗性（adult plant resistance，APR）（图2），通常与模式触发的免疫（pattern-triggered immunity，PTI）有关[25]。在这一防御层级中，定位于细胞质膜上的模式识别受体（pattern recognition receptors，PRRs）扮演着“哨兵”角色，其监测范围极其广泛：不仅能识别病原体共有的保守分子模式（pathogen-associated molecular patterns/microbe- associated molecular patterns，PAMPs/MAMPs），还能感知寄主细胞壁受损时释放的损伤相关分子模式（damage-associated molecular patterns，DAMPs）以及调节免疫平衡的植物细胞因子（phytocytokines）。这种细胞表面感知机制使寄主能够在病原菌建立稳定侵染之前启动防御反应，因而对活体营养型和死体营养型病原菌均具有重要抗性贡献[20]。另外，部分数量抗性还表现出显著的耐病性（disease tolerance）或慢病性（slow-diseasing）特点，通常不诱发剧烈的细胞死亡，而是通过延缓侵染进程、限制菌丝扩展或降低孢子产量来发挥作用，由于不产生强烈的选择压力，此类非特异性的防御策略能显著减缓病原菌毒性的演化速度[25-26]。这种由PTI介导的基础防御与耐病性机制共同构成了小麦实现持久性和广谱抗性的重要基础。
目前，小麦抗性基因的发掘已取得显著进展，尤其在锈病和白粉病抗病基因领域，相关位点已分别编目至Lr85、Yr88、Sr67和Pm70[27]；相比之下，针对赤霉病、茎基腐病等死体营养型病害正式命名的位点均少于20个，但暂时命名的基因和QTL数量极其庞大。为解决历史命名不规范引发的数据整合障碍问题，国际小麦研究界近年更新了基因命名指南[28]，推动抗病基因信息的统一命名与标准化管理，为抗性资源的系统梳理、跨研究比较以及不同防御机制的整合利用提供了重要支撑。除寄主抗性外，非寄主抗性作为植物界最稳定且广谱的免疫形式，虽在目前育种实践中直接利用难度较大，但其所体现的广谱、持久防御机制仍为突破现有抗性易失效问题提供了重要启示。通过深度解析各类抗性资源的分子作用机制，并科学整合不同类型的抗病基因，将为构建小麦全生育期持久抗性屏障奠定坚实基础。
3 小麦抗病基因的发掘、结构多样性及其分子调控机制
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过去20年间，植物免疫研究已从单一基因的克隆转向对免疫复合体结构及其介导的复杂信号网络的系统解构。小麦在与病原菌长期的协同进化过程中，针对活体营养型病原菌的效应子介导侵染和死体营养型病原菌的毒素诱导侵染，演化出具有不同结构特征和作用方式的抗病因子，形成了多样化的免疫调控机制。
3.1 已克隆抗病基因的分布特征与蛋白质结构多样性
在基因组学与克隆技术的推动下，小麦抗病基因的发掘已进入快速积累阶段。针对上述病害，目前已有78个抗病位点、共计101个抗病基因被成功克隆（表1），为抗病育种提供了丰富的遗传资源[14]。从抗性谱分布来看，约85%的基因针对锈病（条锈病16个、叶锈病13个、秆锈病18个）及白粉病（24个）等活体营养型病原菌；而针对赤霉病、叶枯病等死体营养型病原菌的基因仅6个；此外，有3个基因表现出跨病害的广谱持久抗性。在遗传来源方面，已克隆位点在A、B、D亚基因组间分布分别为19、24和21个，其等位变异虽主要源自初级和次级基因库（普通小麦及其祖先供体），但来自三级基因库（野生近缘种）的贡献亦不容忽视，目前已从黑麦（R基因组）、簇毛麦（V基因组）、长穗偃麦草（E基因组）及沙融山羊草（S基因组）中分离出15个具有重要价值的外源基因，极大地拓宽了普通小麦的抗性遗传基础。
根据蛋白质结构特征，这些抗病基因可系统划分为三大功能类群。首先是数量最庞大、研究最为深入的NLR类受体（共52个），其典型结构由NB-ARC结构域与C端LRR识别域组成，通过监测病原菌效应子诱发超敏反应（HR），这类基因介导了典型的针对活体营养型病原菌的小种特异性抗性，是目前抗病育种中应用最广泛的一类抗性基因。其次是激酶及其融合蛋白类（共15个），涵盖了受体样激酶（RLK）、胞质激酶（RLCK）及激酶融合蛋白（KFP），此类基因在应对不同营养模式的病原菌时表现出功能多样性，激酶类受体多参与细胞表面免疫信号的感知与传递，如针对死体营养型病原菌的Stb15和Stb16[29-30]；而激酶融合蛋白则通过结构重组演化出更复杂的监控机制，如介导广谱抗性的Yr15、Yr36、Lr9及Pm24等[31-34]，此类基因通常表现出比NLR更宽的抗性谱及更优的持久性。最后是非经典转运蛋白与多效抗性基因，以Lr34/Yr18（ABC转运蛋白）和Lr67/Yr46（己糖转运蛋白）为典型[35-36]，它们不依赖经典的效应子识别通路，而是通过干扰寄主代谢流或生理平衡来抑制病原菌扩展，其特征是多病害兼抗、非小种特异性且抗性持久。这些不同类群的基因并非孤立存在，最新的研究观点认为，PTI与ETI之间存在深度协同，二者通过复杂的信号网络交互，共同决定了寄主抗性的强度与广度[37-38]。
3.2 NLR抗病小体组装、等位分化与协同识别机制
小麦针对活体营养型病原菌的特异性免疫主要由CC-NLR（CNL）类蛋白介导。当前，这类受体不再被视为孤立的生化开关，而是能够以抗病小体、成对NLR及NLR协同网络等不同组织形式参与免疫调控，其功能机制也由单一识别逐步拓展至更复杂的信号激活过程。
3.2.1 抗病小体的组装与免疫激活
在感知病原菌效应子后，NLR蛋白通过构象重组发生寡聚化，这是启动免疫反应的物理基础。例如，小麦Sr35在识别效应子AvrSr35后组装成五聚体抗病小体，其N端α-螺旋通过翻转插入细胞膜形成阳离子通道，触发超敏反应[39]。这一过程揭示了小麦族CNL类受体启动免疫的保守物理生化机制。此外，最新研究发现，小麦NLR还可组装形成八聚体抗病小体。与此前报道的五聚体CNL抗病小体不同，该八聚体结构呈现独特的漏斗状构架，其CC结构域位于LRR结构域相对一侧，形成有别于经典CNL抗病小体的结构排布方式，进一步拓展了植物NLR抗病小体已知的组装模式和激活机制认知[40]。值得注意的是，NLR介导的HR反应在应对不同营养类型病原菌时表现出明显的功能差异。在抵抗活体营养菌时，HR能够有效限制病原扩展；但在应对死体营养菌时，部分NLR则可能因识别病原菌产生的寄主特异性毒素而“欺骗性”触发HR。由于死体营养型病原菌依赖坏死组织获取营养，这种防御反应反而促进其侵染，形成典型的“反向基因对基因”互作模式。在此情境下，NLR蛋白在功能上转变为“敏感性基因（susceptibility gene，S）”，而有效的防御则需依赖于对细胞死亡的精细抑制及细胞壁加固等非坏死性防御路径（图3）[20,41]。这一现象表明，针对死体营养型病害的改良策略不能简单依赖免疫激活，而应通过基因编辑精准失活或弱化特定的NLR敏感因子，或开发能够屏蔽寄主特异性毒素干扰的小分子拮抗剂，从而实现由感病反应向抗病反应的定向转变，是未来精准育种与绿色防控的重要靶标。
3.2.2 多倍化驱动的等位变异与抗性谱演化
多倍化过程为小麦族NLR基因的进化提供了丰富的遗传素材，促使大量具有新识别特异性的等位基因产生。例如，普通小麦的Pm3位点与黑麦的Pm8、Pm17位点互为同源基因，但在长期的演化中获得了识别不同白粉菌小种的能力[42-43]。更为典型的是，单个NLR基因的不同等位变异可实现抗性特异性的显著分化：如抗叶锈病与条锈病基因Lr13/Yr27（氨基酸差异仅29个）[44-46]、抗白粉病与条锈病基因Pm5/Yr6（差异60个）[47-49]、以及抗秆锈病与白粉病基因Sr22/Pm37（差异54个）（图3）[50-51]。这些案例表明，少数关键氨基酸变异即可重塑NLR受体的病原识别特异性，从而实现抗性谱转换。此外，源自山羊草的Yr87/Lr85展示了另一种广谱抗性形成机制，其LRR重复序列由独特的“双马蹄形”结构组成，可能通过识别保守效应子（或其靶标）来同时抑制叶锈菌和条锈菌的发育[52]。上述研究表明，NLR受体的识别特异性具有较强的可塑性，为抗病受体改造、抗性谱拓展以及新型抗病资源创制提供了重要理论依据。基于结构生物学解析关键识别位点，并结合碱基编辑或先导编辑等精准编辑技术，有望实现感病等位基因的定向优化，加速新型抗病种质的创制。
3.2.3 成对NLRs模式与整合结构域（IDs）的进化演变
小麦中许多NLRs并非单独发挥作用，而是以“传感器-辅助者”或“传感器-执行者”成对的形式协同工作（图3）[53]。研究发现，Pm5（Pm5e和RXL）、Pm26（TdCNL1和TdCNL5）、Pm68及其等位变异（PmWR183-NLR1/2）、Yr6（Yr6RL和Yr6CNL）以及YrTD121/Yr84（TdNLR1/2）等位位点，均由一对功能上紧密耦合的NLRs组成[48-49,54-60]。此外，部分NLRs通过基因融合演化出了特殊的整合结构域（IDs），这些结构域通常来源于病原菌效应子的寄主作用靶标，可参与效应子识别过程。目前在小麦族中已发现整合有锌指BED结构域、锚蛋白重复（ANK）结构域和WRKY转录因子结构域等多种ID的NLR蛋白。例如，小麦的Yr5a/b/x和Yr7均含有BED结构域[61-62]，而YrU1则同时整合了ANK和WRKY两种结构域，极大地扩展了植物应对病原菌攻击的监测范围[63]。NLR-IDs的形成机制表明，植物能够通过整合新的功能结构域不断扩展病原感知能力，这为人工设计具有新识别特异性的抗病受体提供了重要启示。通过定向引入效应子靶标结构域或优化现有识别模块，可以创制出针对新兴毒性小种的广谱监控器，这为小麦抗病基因库的人工扩容与抗性持久性的动态演化提供了创新的技术路线。
3.3 激酶类受体与融合蛋白介导的免疫调控
小麦针对死体营养型病原菌的抗性主要由模式识别受体PRRs介导。例如，细胞壁相关激酶（WAKs）Stb6与G型凝集素受体激酶（LecRK）Stb15能够分别特异性识别壳针孢叶斑病菌的效应子AvrStb6和AvrStb15来触发抗病性[30,64]；而Stb16q编码一种富含半胱氨酸的受体样激酶（CRKs），能抑制叶枯病菌的气孔侵染，显著减少病原菌早期的胞间增殖[29]。PRRs的识别同样具有复杂的两面性，病原菌常通过劫持寄主的免疫反应来实现侵染，如编码WAK的Snn1在识别颖枯病菌效应子SnTox1后反而会诱导细胞死亡，从而有利于病原菌的定殖，表现为典型的感病特征[65]。相比于单一抗病位点的解析，提升基础免疫的广谱性更具育种价值，以LysM受体介导的几丁质识别系统为例，将小麦近缘属种簇毛麦的几丁质受体基因CERK1-V导入普通小麦后，其编码蛋白能够与内源受体TaCEBiP形成免疫复合体，通过激活MAPK/CPK磷酸化、活性氧猝发及水杨酸与茉莉酸等激素信号通路，显著协同提升小麦对白粉病、条锈病及赤霉病的抗性（图3）[66]。这一研究表明通过引入外源或优化关键模式识别受体，可增强小麦对多种病原菌的基础免疫能力，为广谱抗性改良提供新的遗传资源。
激酶融合蛋白是近年来在麦类作物基因组中发现的一类重要非典型R基因，它们通过激酶结构域（KD）与其他功能域的模块化整合，突破了传统免疫受体相对固定的结构组成方式，从而形成更加多样化的病原感知和信号转导机制[67-68]。由于同时具备病原识别和信号传递相关功能，部分激酶融合蛋白兼具PTI与ETI特征，因此被认为是植物免疫系统中具有重要研究价值的一类受体蛋白。
3.3.1 串联激酶蛋白（TKP）的识别特异性与复合免疫模块
这类蛋白通常含有两个串联的激酶（或假激酶）结构域，已成为小麦抗病研究的热点。研究证实，抗麦瘟病蛋白RWT4通过其N端激酶部分重复（KDup）的关键氨基酸（V124/R125）和假激酶结构域（PKD）直接结合效应子AvrPWT4，进而解除分子内抑制，激活自身激酶活性与下游细胞死亡信号，揭示了TKP可直接参与病原菌效应子的识别过程[69]。同时，TKP还能通过跨家族协作形成“激酶-NLR”复合防御模块（图3）。例如，抗白粉病蛋白Pm24（WTK3）通过假激酶与Kin I结构域感知效应子后，利用Kin II结构域衔接NLR蛋白WTN1，组装成促进钙离子（Ca2+）内流的复合物以激活超敏反应[70]；抗秆锈病蛋白Sr62TK的Kinase 1结构域在静息态下抑制Kinase 2，当效应子AvrSr62与其Kinase 1竞争性结合后释放Kinase 2，使其暴露互作界面并激活下游Sr62NLR触发免疫[71]。表明部分TKP并非独立发挥作用，而是与NLR受体形成功能耦联的识别与信号转导体系。这种机制与拟南芥中经典的ZAR1-RKS1-PBL2三组分免疫复合体具有较高相似性[72]。这些发现表明激酶蛋白不仅能够作为病原感知元件，还能够参与免疫信号的传递与放大过程。从应用角度看，这类受体体系具有较强的可塑性。通过对效应子识别相关结构域进行定向改造，理论上可拓展其病原识别范围，为新型抗病受体的设计与创制提供重要参考。
3.3.2 激酶-整合域融合蛋白（KD-ID）的重组与防御功能特征
此类蛋白通过KD域与不同整合域（ID）的融合实现了功能的极大拓展：如抗秆锈病蛋白Sr43集成了两个未知功能的DUF结构域；抗白粉病蛋白Pm13则由N端混合谱系激酶假激酶域（MLKL_NTD）与C端丝氨酸/苏氨酸真激酶（STK）构成，与动植物经典MLKL假激酶执行细胞死亡的机制不同，Pm13形成了由激酶结构域N端α螺旋直接介导细胞死亡的作用机制[73-74]；抗白粉病蛋白Pm4b通过激酶域与多个C2结构域融合，利用选择性剪接在内质网上形成异源复合物，以此实现对白粉病抗性的精准调控[75]。最新研究揭示了病原菌针对Pm4的进化逃逸策略：白粉菌效应子特定变异体SvrPm4（即AvrPm1a）能作为强效抑制子阻断AvrPm4与Pm4触发的免疫反应，但该变异体仍可被Pm1a识别，反映出病原菌毒力演化与寄主免疫监控之间持续的协同进化过程[76]。然而，KD-ID介导的抗性并不总是表现为正向防御效应。部分融合受体如Tsn1和Snn3-D1，在分别识别褐斑叶枯病菌效应子ToxA或颖枯病菌效应子SnTox3后，会被死体营养型病原菌利用，从而由“防御哨兵”异化为病原菌入侵的易感靶标[41]；相比之下，抗条锈病蛋白Yr36（WKS1）则展现出激酶与START结构域融合后的独特防御机制：其进入叶绿体后通过磷酸化PsbO削弱光合作用并促进H2O2积累，同时磷酸化tAPX抑制其清除活性氧的能力，从而诱导局部活性氧积累和细胞死亡，最终限制病原菌扩展[77-78]。这些研究表明整合结构域的获得不仅能够拓展受体蛋白的功能边界，也可能导致防御反应在不同病原侵染背景下产生差异化结果。这种结构与功能的多样化演化，为理解植物免疫系统的适应性创新提供了重要案例。同时，不同功能结构域之间的可组合性也为抗病受体改造和新型抗病资源创制提供了理论依据，是未来受体工程和精准抗性改良的重要靶标。
3.3.3 等位变异驱动的广谱兼抗特性
串联激酶蛋白与激酶融合蛋白家族展现出极强的抗性谱演化潜力，其单倍型之间的少量序列差异即可导致病原识别特异性的改变。例如，Pm24与Rwt4互为等位变异，除对白粉病具有抗性外，还能够介导对麦瘟病的抗性[70,79]；与之类似，起源于Pm4家族的Rmg7/Rmg8同样展现出对白粉病与麦瘟病的多重防御能力[80-81]；此外，抗叶锈病基因Lr9与抗白粉病基因Pm57编码的蛋白同源性高达88%，且均包含融合vWA结构域，分别赋予小麦对不同病害的抗性[34,82]。这种“一因多抗”的特性揭示了激酶类抗病基因能够通过有限的序列变异实现病原识别范围的调整与扩展，从而形成多样化的抗性谱。这一现象反映了植物在长期协同进化过程中对病原菌选择压力的持续适应，也揭示了等位变异在抗病功能分化中的重要作用。从育种利用角度看，解析决定抗病基因抗性特异性的关键变异位点，通过精准编辑手段对感病品种进行内源性重塑，定向优化现有等位基因，快速创制具有更优抗性谱的种质资源，可为小麦广谱持久抗性的培育提供新的途径。
3.4 非经典抗性基因介导的生理稳态重构与广谱防御机制
不同于以直接识别病原菌效应子为主要特征的典型免疫，小麦中还存在一类通过调控寄主自身生理状态实现广谱持久抗性的非经典基因。这类基因通常不含典型抗病结构域，而是通过影响寄主的能量代谢、离子稳态以及蛋白质动态行为等过程，增强植株对病原菌侵染的适应与防御能力。
3.4.1 转运蛋白驱动的能量代谢与营养免疫
小麦通过改变糖类及脂质的分配权衡，实现了对病原菌的“养分封锁”（图3）。广谱持久抗性基因Yr18/Lr34编码一种ABC转运蛋白，其抗病等位变异通过调控磷脂代谢如促进磷脂酸向内、磷脂酰丝氨酸向外转运来改变胞膜状态，并利用其对脱落酸（ABA）的转运能力增强生物胁迫响应[35,83-84]；Yr46/Lr67则编码一种己糖转运蛋白，抗病突变导致其丧失葡萄糖转运活性并产生由阴离子主导的内向电流，从而限制病原菌对寄主糖源的利用并抑制侵染过程[36,85-86]。另外，含有脂质结合START结构域的蛋白可能是广谱抗性演化的另一关键模块。广谱抗病基因Yr36与YrKB（TaEDR2-B）均以此为感应枢纽，虽与拟南芥感病基因AtEDR2同源，但可能通过结构域重组实现了从免疫抑制向抗病激活的功能分化[62]。START域作为脂质感应开关，可通过特异性结合磷脂诱导蛋白变构，促进下游激酶或转录因子的二聚化与激活，将胞内脂质波动转化为免疫输出。这种潜在的配体感应机制解释了YrKB兼顾广谱抗性与产量稳态（no yield penalty）的特性，为解析脂质代谢流权衡抗性与发育提供了重要分子模型。
3.4.2 离子通道介导的气孔免疫与跨细胞协同防御
气孔作为病原菌入侵的首要门户，其免疫动态由精准的离子流调控。最新研究揭示，含锚蛋白重复序列（ANK）的跨膜蛋白Lr14a本质上是一种功能性Ca2+通道（图3）[87]。被病原菌诱导后，Lr14a在叶肉细胞中激活，通过一种创新的非细胞自主信号转导途径远程操控保卫细胞发生HR。这种跨细胞信号传递机制实现了侵染部位与气孔防御反应之间的协同调控，使植株能够在避免大面积组织坏死的情况下有效限制病原菌扩展[88]。
3.4.3 毒素中和与相分离驱动的蛋白枢纽调控
针对赤霉病等释放毒素的病害，小麦演化出了直接解毒与精细调控大分子凝聚态两种不同的防御策略（图3）。一方面，Fhb7编码的谷胱甘肽S-转移酶（GST）能通过催化DON毒素与谷胱甘肽结合实现直接解毒[89]。另一方面，针对核心位点Fhb1的最新研究揭示，其关键基因编码蛋白TaHRC通过“液-液相分离”形成的蛋白枢纽状态决定了植株的抗感特性。感病型TaHRC-S在DON毒素诱导下易发生相分离并驱动招募剪切体组分（如TaSR45a）等7个成员形成凝聚态枢纽（hub），通过改变免疫相关基因的可变剪接并促进DON介导的细胞死亡导致感病；而抗病型TaHRC-R因N端关键半胱氨酸变异导致构象改变，不仅自身对DON诱导不敏感，还能抑制复合体成员的相分离，使其维持在非凝聚态，从而保障了免疫信号的正常传导并赋予小麦强效抗性[90]。这种通过调控生物大分子凝聚状态来应答病原菌侵染的新机制，为解析小麦多病害非特异性抗性提供了全新的物理生化视角。
这类非经典基因虽然缺乏特定抗病结构域，且常伴随细胞衰老加速等生理代价，但其核心优势在于靶向寄主自身基础生理过程，有效规避了病原菌通过效应子变异而逃避识别的缺陷[91]。这些机制的解析，标志着小麦防御研究正从传统的受体识别视角转向细胞微环境重塑的新维度。这不仅为通过微调转运蛋白基质特异性或利用小分子干预蛋白相分离来实现内生性广谱抗性改良奠定了基础，更为开发精准靶向养分流向、离子稳态及物理状态的新型绿色防控技术开辟了路径。
3.5 病原菌对寄主生理稳态的精准劫持
从生物演化视角看，感病并非植物主动演化形成的功能，而是病原菌与寄主协同进化过程中，病原菌利用植物基础生理通路引发的结果。这种易感性本质来源于植物正常生理过程中的薄弱环节：一方面源于寄主自身的负向免疫调控机制，用以防止免疫持续激活造成大量能量损耗；另一方面则是由于某些执行基础代谢或发育功能的基因（S）被病原菌效应子精准劫持[92]。因此，通过失活这些易感相关位点，本质上是阻断了病原菌利用寄主资源的通路，从而获得持久且广谱的隐性抗性。
3.5.1 核心防御稳态的负调控机制利用
典型代表是MLO。MLO蛋白定位于质膜，通过抑制侵染位点细胞壁的活性氧爆发来维持胞间稳态，但这客观上为白粉菌的侵入提供了便利[93-95]。小麦TaMLO突变虽能赋予广谱抗性，但也易诱发早衰[96]。小麦Tamlo-R32通过大片段缺失引发的染色体结构改变，可诱导上游基因TaTMT3表达，从而在维持高抗性的同时补偿MLO缺失带来的产量损失，实现了防御与生长平衡的精准重构[97]。
3.5.2 基础信号通路的效应子劫持与信号放大
病原菌常将寄主正常的信号传递蛋白转化为辅助侵染的内线。例如，条锈菌效应子PsSpg1通过劫持小麦激酶TaPsIPK1，增强其入核磷酸化转录因子TaCBF1d的能力，从而抑制防御基因表达并反馈放大自身的感病信号[98]。这种机制揭示了病原菌如何通过调控寄主内源信号流，将植物的正常代谢响应重构为利于侵染的感病状态。
4 小麦抗病性改良的多元遗传策略
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随着抗病基因资源的持续积累以及植物免疫机制研究的不断深入，小麦抗病育种正由传统的经验选择逐步向精准设计育种发展。近年来，大量抗病基因、数量性状位点（QTL）及其作用机制被解析，为抗性资源的高效利用提供了重要基础。与此同时，病原菌群体持续演化以及多病害复合发生趋势，也对抗性改良提出了更高要求。因此，如何实现抗性资源的科学配置与精准利用，培育兼具广谱性、持久性和农艺适应性的新品种，已成为当前小麦抗病育种的重要研究方向。基于此，本章重点围绕抗性资源利用与遗传改良策略展开讨论，包括基于全景抗性图谱的抗病位点精准布局与聚合利用、野生近缘种优异抗性资源的发掘与导入、基因编辑与免疫受体工程驱动的抗性定向改良，以及基因芯片和数字化平台支撑的精准设计育种等方面，为小麦广谱、持久抗性材料的创制提供参考。
4.1 遗传全景图谱构建与基因精准聚合策略
针对活体寄生病害，小麦抗病育种长期面临病原菌毒性演化快、品种抗性易丧失以及研究信息碎片化等多重挑战。由于不同抗病位点的遗传效应、利用价值及区域适应性缺乏系统整合，抗病资源的利用往往呈现分散化和经验化特点。因此，通过全球尺度的资源整合，构建一张可检索、可验证、可预测的基因组“导航图”，是实现从经验育种向精准设计育种跨越的重要基础。笔者团队通过整合全球近1.5万份种质资源及4.7万个条锈病抗性表型数据，利用全基因组关联荟萃分析，绘制出包含431个QTL位点的小麦抗条锈病全景图[62]。该图谱不仅厘定了如Yr18、Yr29等经典抗性位点的全球分布，更发掘出172个具有潜在应用价值的新位点。通过对百年间抗性等位基因频率的时空评估，还原了百年来寄主-病原菌的共进化轨迹，揭示了苗期抗性基因的动态更替过程，并捕捉到欧洲Pst S系列小种及中国V26毒性群体扩散对全球小麦抗性格局的影响。同时，空间维度分析明确了全球广适性抗性热点区域以及具有显著地域特征的选择位点，揭示了不同麦区骨干亲本对抗性位点频率形成的差异化影响，为抗病基因的科学轮换与区域化精准布局提供了决策依据。除单一病害外，相关研究进一步将分析范围拓展至条锈病、叶锈病、秆锈病、白粉病和赤霉病等主要病害。通过对2 484个QTL进行元分析，整合获得732个独立QTL，其中317个为介导多病害抗性的位点（未发表）。需要指出的是，全景导航图并非旨在将数百个基因或QTL同时导入育种材料，而是为不同生态区和育种目标提供科学的位点筛选依据。通过厘清不同抗病位点的遗传效应、地域适应性及互作关系，可建立“核心主效基因+区域适应性位点+广谱持久抗性位点”的分层配置体系。例如，在条锈病高发区可优先聚合广谱持久抗性基因与当地主导毒性小种对应的主效基因；而在多病害复发麦区，则可重点利用具有多病害抗性的共享位点进行协同聚合。因此，全景导航图的价值不仅体现在抗病新位点的发掘，更体现在为不同生态区域制定差异化抗病基因组合方案提供依据。通过明确“哪些基因值得用、哪些基因适合在何处使用以及哪些基因适合联合利用”，能够为抗病基因的精准布局和科学轮换提供支撑。
4.2 深度挖掘野生近缘种抗性资源以拓宽免疫遗传基础
小麦族野生近缘植物包含30个属、400多个种，是栽培小麦应对生物胁迫的重要遗传资源库。受驯化过程中遗传瓶颈的影响，栽培小麦的免疫多样性显著收缩，而野生近缘种在长期自然演化中积累了丰富的抗病等位基因。据不完全统计，已从野生种中鉴定并转移了200余个抗锈病、抗白粉病等关键基因到栽培小麦[99]，因此，持续发掘和利用野生近缘种中的优异抗病资源，已成为拓展小麦抗性遗传基础的重要途径。近年来，基因组学技术的发展显著提升了野生种优异等位变异的发掘效率。作为栽培小麦的直接祖先，野生二粒小麦（Triticum dicoccoides）因与普通小麦杂交亲和性好、且兼具高蛋白与多抗特性，成为抗病资源利用最广泛的野生供体之一，已从中克隆并利用了Yr15、Yr36、Pm26、Pm36等[100]。同时，针对D基因组供体山羊草（Aegilops tauschii）的大规模全基因组重测序，进一步加深了对其遗传多样性和演化过程的认识。通过对46个粗山羊草基因组的单倍型分析，研究者已能精准区分复杂位点上的旁系同源拷贝，实现了对Sr66、Lr39等新型抗病基因的精准定位与克隆[101-102]。抗病基因挖掘策略正从单纯的表型驱动转向高通量表达辅助驱动。传统观点认为NLR类抗病基因主要受诱导表达，但研究发现大量功能性NLR在健康植物组织中即维持较高水平表达。基于此，建立的集高表达筛选、高通量转化、大规模表型鉴定为一体的流程，已成功从小麦近缘禾本科植物中构建了包含995个候选NLR的转基因文库，并快速鉴定出31个新型抗锈病基因[103]。随着野生种基因组资源的不断完善以及功能验证效率的持续提升，抗病基因发掘正由随机筛选逐步转向目标导向的精准挖掘。这不仅提高了优异抗病资源的利用效率，也为持续拓展小麦抗病基因库提供了重要支撑。
4.3 基因组编辑驱动的受体定向进化与感病基因定向敲除
基于对寄主易感性基础及受体激活机制的不断深入认识，抗性改良已逐步从传统抗源筛选拓展至对关键遗传元件的精准改造。针对感病基因（S）这一病原菌侵染所必需的寄主组分，通过精准干预其介导的入侵门户、代谢补给或免疫负调控等功能，可赋予小麦更为持久广谱的抗性。典型案例是在小麦品种科农199中对小麦MLO的工程化改造以及在品种百农207、扬麦158等敲除感病基因TaPsIPK1，所得突变体表现出广谱抗性，且分蘖、千粒重等农艺性状不受影响，实现了抗病性与农艺性状的协调改良[97-98]。与此同时，针对抗性受体本身的精准改造正成为抗病育种的重要研究方向。一方面，利用碱基编辑技术可对感病与抗性等位基因间的关键氨基酸位点（如Lr30、Sr13等）进行定向转换，或通过启动子编辑恢复被抑制的抗性基因表达（如Pm41b）[104-106]；另一方面，随着结构生物学的发展，研究者开始尝试依据受体结构信息开展理性设计，例如以Sr33为骨架重塑LRR结构域的识别特异性[107]，或利用Pm5/Yr6等具有识别可塑性的天然等位变异构建新型免疫受体。相比传统依赖自然变异的育种方式，这类策略能够更有针对性地拓展受体识别范围，提高对新兴毒性小种的响应能力。从感病基因精准敲除到抗性受体定向优化，基因组编辑正在为小麦抗病改良提供更加灵活的遗传操作手段。随着受体识别机制和病原菌致病规律研究的不断深入，基于理性设计的抗性创制将成为未来培育广谱、持久抗病品种的重要途径。
4.4 基因芯片与数字化平台驱动的抗性精准设计育种
随着种质资源研究从表型鉴定逐步进入全基因组解析阶段，小麦抗病育种正在由经验选择向数据驱动的精准设计育种转变。在这一过程中，高通量液相捕获芯片（如GenoBaits® WheatSNP系列）与数字化育种平台的结合，为抗病基因的发掘、跟踪与利用提供了重要技术支撑。通过将基因型数据、表型数据与育种信息进行系统整合，研究者能够更加高效地解析抗性遗传基础，并为抗病基因的组合利用提供依据。高密度背景芯片显著提高了抗病位点的定位效率与解析精度。借助GBW16K等覆盖全基因组的液相背景芯片，研究者能够快速构建遗传图谱并开展QTL精细定位[108]。这一技术已成功应用于Yr18、Yr27、Yr30等经典位点的快速鉴定，并辅助发掘了QYr.nwafu-2BL.4等新抗病位点。针对远缘杂交育种中外源片段难以精确鉴定的问题，开发的小麦-长穗偃麦草、小麦-黑麦等系列专用芯片实现了对外源染色体断点位置及拷贝数的精准追踪，显著提高了外源抗病资源导入与评价的效率[109]。与此同时，数字化平台与前景选择芯片的结合进一步提升了抗病基因的利用效率。通过集成抗病、产量等核心目标性状的GBW0.1K前景芯片，研究者实现了对我国主栽品种抗性背景的系统解析，明确了Yr29、Yr30等重要基因的分布格局[110]。在此基础上，构建的小麦锈病抗性位点全景图数据库（LWRR）整合了全球2 191份种质资源、431个QTL及9 276个候选基因信息，并提供在线查询与可视化分析功能。该平台能够帮助研究者快速获取目标位点的分布特征、候选基因信息及区域适应性规律，为抗病基因的区域化配置和科学利用提供参考[111]。在育种实践中，芯片检测、算法导航与数字化平台的协同应用，正在推动抗病育种由经验判断向数据辅助决策转变。例如，通过联合应用“GBW16K背景+GBW0.1K前景”芯片，可快速解析西农302、西农877等新品种（系）的基因组组成及抗病位点构成[112-113]，为后续育种决策提供依据。
无独有偶，为解决优异资源向育种背景转化效率较低的问题，张学勇团队构建了回交-互交-巢式关联作图群体（AB-NAMIC）[114]。该群体以主栽品种为受体、核心种质为供体，通过回交与亚群互交实现骨干亲本遗传背景与优异资源的有效融合。与自然群体相比，AB-NAMIC显著提高了主效基因及稀有有利等位变异的频率，使TaSWEET6-7B等中低效应位点得以稳定检测，并提升了复杂农艺性状和抗病性状的遗传解析能力。依托该群体建立的WheatGAB数据库进一步整合了1 428个有利等位变异，其中包括300个与锈病和白粉病等相关的抗性位点。利用该群体训练的全基因组选择（GS）模型，其预测精度明显优于自然群体。这一研究表明，通过遗传资源创新、群体构建与数字化分析平台的结合，能够有效提升优异等位变异的发掘与利用效率，为多性状精准聚合探索出数字化育种新途径。
5 小麦抗性创新利用展望
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面对全球气候变化背景下病原菌传播范围扩大、毒性结构持续演化以及多病害复合发生等挑战，传统依赖表型鉴定和回交转育的抗病育种模式正面临新的考验。随着基因组学、人工智能（AI）、合成生物学和计算生物学的快速发展，小麦抗病研究正在从抗性资源发掘逐步延伸至抗性系统的预测、设计与重构。未来，小麦抗病改良将进一步实现从经验驱动向数据驱动、从被动响应向前瞻设计的转变。
5.1 基于基因组定点操作的抗性模块化组配
针对复杂农艺性状改良过程中存在的连锁累赘及多基因聚合效率受限等问题，未来借助基因组定点编辑与大片段精准整合技术，实现抗性位点的模块化组配。通过在染色体上建立稳定表达且对植株生长影响较小的“基因组安全港”，结合先导编辑及定点整合技术，打造类似“USB接口”的即插即用型免疫模块的育种平台。在此基础上，结合快速育种和高通量表型分析技术，进一步缩短抗性材料创制与评价周期，提高多抗材料的选育效率。未来，围绕特定生态区病原菌群体结构开展针对性的抗性位点组合设计，可能成为区域化精准抗病育种的重要发展方向（图4）。
5.2 构建免疫数字孪生系统引导的预测性抗病设计
未来抗性发掘将不再依赖于随机的自然突变，而是迈向全谱系的智能设计。通过整合覆盖禾本科广谱等位基因的泛R图谱与病原菌泛效应因子组，利用大语言模型（LLM）与蛋白质结构预测技术（如AlphaFold）构建寄主-病原菌互作的数字模型。这一“免疫数字孪生”系统允许研究者在计算机中模拟效应子的演化轨迹及识别残基的稳定性，从而在田间变异发生前完成抗病受体的定向进化设计。这种从灾后响应向先验设计的转变，是提升抗病持久性的核心战略储备（图4）。
5.3 通用受体设计与多界免疫生态
利用理性设计突破自然选择瓶颈，是构建超越单一遗传背景多维防御体系的核心。针对活体与死体营养型病原菌在免疫响应上的差异，未来小麦抗病研究的重要方向之一是探索防御激活与细胞死亡之间的相对解耦机制。对于活体营养型病原菌，细胞死亡有助于限制病原扩展；而对于死体营养型病原菌，过度的细胞死亡则可能被病原菌利用。如何在维持免疫强度的同时降低细胞死亡带来的负面效应，是提升广谱抗性的重要科学问题。随着结构生物学、系统发育分析及人工智能辅助蛋白设计的发展，研究者开始尝试从不同病原菌中鉴定保守效应子靶点，并据此优化受体识别能力。通过对NLR整合结构域、RLK激酶结构域等关键区域进行定向改造，设计出通用免疫受体，拓展天然受体的识别范围，并减少病原菌利用宿主敏感因子诱导感病的风险。这类研究为构建兼具广谱性与持久性的免疫受体提供了新的思路。与此同时，植物抗病能力的研究视角正在由寄主基因组逐步扩展至更广泛的生物互作体系。越来越多的研究表明，植物不仅依赖自身免疫系统抵御病原菌侵染，还能够通过分泌香豆素等代谢物调节根际微生物群落组成，间接增强抗病能力。尤其在应对死体营养型病原菌时，益生微生物可通过竞争营养资源、调节活性氧稳态及诱导系统性抗性等方式参与病害抑制，从而形成寄主免疫的重要补充。未来有必要从寄主-微生物-病原菌三方互作的角度重新认识植物抗病机制。结合多界GWAS、宏基因组学和合成群落构建等技术，解析不同生物组分之间的互作规律，并筛选具有稳定促抗功能的核心微生物群落，将有助于构建兼顾遗传抗性与生态稳定性的综合防御体系。这种将寄主遗传改良与微生物生态调控相结合的策略，可能为应对病原菌持续演化和多病害复合发生提供新的解决途径（图4）。
综上所述，随着小麦抗病基因资源的持续发掘、免疫调控机制的深入解析以及基因组编辑、人工智能和合成生物学等新技术的发展，小麦抗病研究正逐步由抗性资源利用迈向免疫系统的理性设计。未来，基于抗病基因全景解析、受体功能优化、多病害协同防御以及寄主-微生物互作调控等方向的持续探索，将进一步丰富对植物免疫体系的认识，并为广谱、持久抗性的构建提供新的理论依据和技术路径。
从更广泛的视角来看，小麦抗病改良不仅是抗性基因的聚合与利用，更是对寄主与病原菌长期协同进化规律的认知与重构。随着多学科交叉融合的不断深入，抗病育种有望逐步实现由经验选择向精准设计转变，由单一基因改良向系统性防御优化拓展。如何在提升抗性的同时兼顾产量、品质与生态适应性，仍将是未来研究需要持续关注的重要命题。相关理论与技术的发展，将为复杂病害环境下小麦稳产增产及农业可持续发展提供重要支撑。
基金
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  • 国家自然科学基金(32272088)
  • 国家自然科学基金(32225041)
  • 国家自然科学基金(32472103)
  • 国家自然科学基金(32561143296)
  • 国家重点研发计划(2021YFD1401000)
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2026年第59卷第16期
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doi: 10.3864/j.issn.0578-1752.2026.16.007
  • 接收时间：2026-03-22
  • 首发时间：2026-09-03
  • 出版时间：2026-08-16
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  • 收稿日期：2026-03-22
  • 录用日期：2026-06-06
基金
国家自然科学基金(32272088)
国家自然科学基金(32225041)
国家自然科学基金(32472103)
国家自然科学基金(32561143296)
国家重点研发计划(2021YFD1401000)
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    1 西北农林科技大学农学院， 陕西杨凌 712100
    2 西北农林科技大学植物保护学院， 陕西杨凌 712100
    3 作物抗逆与高效生产全国重点实验室， 陕西杨凌 712100

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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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