小麦抗病机制研究进展与抗性创新利用策略

doi:10.3864/j.issn.0578-1752.2026.16.007

摘要/Abstract

摘要：

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。

关键词: 小麦, 抗病基因, 抗/感病机制, 抗病性改良

Abstract:

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.

Key words: 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[J]. 中国农业科学, 2026, 59(16): 3556-3576.

WU JianHui, ZENG QingDong, LIU ShengJie, WANG XiaoJie, HAN DeJun, KANG ZhenSheng. Advances in Mechanisms and Innovative Improvement Strategies for Wheat Disease Resistance[J]. Scientia Agricultura Sinica, 2026, 59(16): 3556-3576.

图/表 5

图1

图2

表1

已克隆的小麦抗病基因"

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	2A^mL	一粒小麦T. monococcum	秆锈病Stem rust (Pgt)
Sr22a/b⁴	CNL	7A^bL; 7A^mL	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	3A^mL	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	5A^mS	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	1S^shS	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	3S^lS	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	6S^lL	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	6S^shS/6S^lS	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)

表1

图3

图4

参考文献 114

[1]	Food and Agriculture Organization of the United Nations (FAO). FAOSTAT. Crops and livestock products[DB/OL]. (2025-12-31) [2026-03-05]. https://www.fao.org/faostat/en/#data/QCL.
[2]	Food and Agriculture Organization of the United Nations (FAO). In brief to the state of the world’s land and water resources for food and agriculture 2025. The potential to produce more and better[DB/OL]. [2026-03-05]. https://doi.org/10.4060/cd7598en.
[3]	Savary S, Willocquet L, Pethybridge S J, Esker P, McRoberts N, Nelson A. The global burden of pathogens and pests on major food crops[J]. Nature Ecology & Evolution, 2019, 3(3): 430-439.
[4]	Bebber D P. Range-expanding pests and pathogens in a warming world[J]. Annual Review of Phytopathology, 2015, 53: 335-356. doi: 10.1146/annurev-phyto-080614-120207 pmid: 26047565
[5]	Chen X. Pathogens which threaten food security: Puccinia striiformis, the wheat stripe rust pathogen[J]. Food Security, 2020, 12(2): 239-251. doi: 10.1007/s12571-020-01016-z
[6]	Kolmer J A, Herman A, Ordoñez M E, German S, Morgounov A, Pretorius Z, Visser B, Anikster Y, Acevedo M. Endemic and panglobal genetic groups, and divergence of host-associated forms in worldwide collections of the wheat leaf rust fungus Puccinia triticina as determined by genotyping by sequencing[J]. Heredity, 2020, 124(3): 397-409. doi: 10.1038/s41437-019-0288-x pmid: 31863032
[7]	Zhang N, Liao Z, Wu S, Nobis M P, Wang J, Wu N. Impact of climate change on wheat security through an alternate host of stripe rust[J]. Food and Energy Security, 2022, 11: e356. doi: 10.1002/fes3.v11.1
[8]	Sotiropoulos A G, Arango-Isaza E, Ban T, Barbieri C, Bourras S, Cowger C, Czembor P C, Ben-David R, Dinoor A, Ellwood S R, Graf J, Hatta K, Helguera M, Sánchez-Martín J, McDonald B A, Morgounov A I, Müller M C, Shamanin V, Shimizu K K, Yoshihira T, et al. Global genomic analyses of wheat powdery mildew reveal association of pathogen spread with historical human migration and trade[J]. Nature Communications, 2022, 13: 4315. doi: 10.1038/s41467-022-31975-0 pmid: 35882860
[9]	Feurtey A, Lorrain C, McDonald M C, Milgate A, Solomon P S, Warren R, Puccetti G, Scalliet G, Torriani S F, Gout L, Marcel T C, Suffert F, Alassimone J, Lipzen A, Yoshinaga Y, Daum C, Barry K, Grigoriev I V, Goodwin S B, Genissel A, et al. A thousand-genome panel retraces the global spread and adaptation of a major fungal crop pathogen[J]. Nature Communications, 2023, 14: 1059. doi: 10.1038/s41467-023-36674-y pmid: 36828814
[10]	康振生, 王晓杰, 赵杰, 汤春蕾, 黄丽丽. 小麦条锈菌致病性及其变异研究进展[J]. 中国农业科学, 2015, 48(17): 3439-3453. DOI: 10.3864/j.issn.0578-1752.2015.17.011.
	Kang Z S, Wang X J, Zhao J, Tang C L, Huang L L. Advances in research of pathogenicity and virulence variation of the wheat stripe rust fungus Puccinia striiformis f. sp. tritici[J]. Scientia Agricultura Sinica, 2015, 48(17): 3439-3453. DOI: 10.3864/j.issn.0578-1752.2015.17.011. (in Chinese)
[11]	Singh R P, Hodson D P, Singh P K, Lan C X, He X Y, Lagudah E S, Juliana P, Ayliffe M, Bhavani S, Saunders D G O, Huerta-Espino J. Challenges to wheat disease resistance and current global strategies[J]. Annual Review of Phytopathology, 2025, 63: 201-224. doi: 10.1146/phyto.2025.63.issue-1
[12]	Pequeno D N, Ferreira T B, Fernandes J M, Singh P K, Pavan W, Sonder K, Robertson R, Krupnik T J, Erenstein O, Asseng S. Production vulnerability to wheat blast disease under climate change[J]. Nature Climate Change, 2024, 14(2): 178-183. doi: 10.1038/s41558-023-01902-2
[13]	韩德俊, 康振生. 中国小麦品种抗条锈病现状及存在问题与对策[J]. 植物保护, 2018, 44(5): 1-12.
	Han D J, Kang Z S. Current status and future strategy in breeding wheat for resistance to stripe rust in China[J]. Plant Protection, 2018, 44(5): 1-12. (in Chinese)
[14]	Li Y, Govta L, Sung Y, Coaker G, Fahima T. The spectrum of diverse disease-resistance genes cloned and characterized in the Triticeae tribe[J]. Annual Review of Phytopathology, 2025, 63: 175-200. doi: 10.1146/phyto.2025.63.issue-1
[15]	Singh P K, Gahtyari N C, Roy C, Roy K K, He X Y, Tembo B, Xu K J, Juliana P, Sonder K, Kabir M R, Chawade A. Wheat blast: A disease spreading by intercontinental jumps and its management strategies[J]. Frontiers in Plant Science, 2021, 12: 710707. doi: 10.3389/fpls.2021.710707
[16]	Saintenac C, Lee W, Cambon F, Rudd J J, King R C, Marande W, Powers S J, Bergès H, Phillips A L, Uauy C, Hammond-Kosack K E, Langin T, Kanyuka K. Wheat receptor-kinase-like protein Stb6 controls gene-for-gene resistance to fungal pathogen Zymoseptoria tritici[J]. Nature Genetics, 2018, 50(3): 368-374. doi: 10.1038/s41588-018-0051-x pmid: 29434355
[17]	McDowell J M. Genomes of obligate plant pathogens reveal adaptations for obligate parasitism[J]. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(22): 8921-8922. doi: 10.1073/pnas.1105802108 pmid: 21576481
[18]	Glazebrook J. Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens[J]. Annual Review of Phytopathology, 2005, 43: 205-227. pmid: 16078883
[19]	Mengiste T. Plant immunity to necrotrophs[J]. Annual Review of Phytopathology, 2012, 50: 267-294. doi: 10.1146/annurev-phyto-081211-172955 pmid: 22726121
[20]	Mengiste T, Liao C. Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens, 20 years later: What has changed?[J]. Annual Review of Phytopathology, 2025, 63: 279-308. doi: 10.1146/phyto.2025.63.issue-1
[21]	Chen Y, Kistler H C, Ma Z. Fusarium graminearum trichothecene mycotoxins: Biosynthesis, regulation, and management[J]. Annual Review of Phytopathology, 2019, 57: 15-39. doi: 10.1146/annurev-phyto-082718-100318 pmid: 30893009
[22]	Biffen R H. Mendel’s laws of inheritance and wheat breeding[J]. The Journal of Agricultural Science, 1905, 1(1): 4-48. doi: 10.1017/S0021859600000137
[23]	Flor H H. Current status of the gene-for-gene concept[J]. Annual Review of Phytopathology, 1971, 9: 275-296. doi: 10.1146/phyto.1971.9.issue-1
[24]	Jones J D G, Dangl J L. The plant immune system[J]. Nature, 2006, 444(7117): 323-329. doi: 10.1038/nature05286
[25]	Niks R E, Qi X, Marcel T C. Quantitative resistance to biotrophic filamentous plant pathogens: Concepts, misconceptions, and mechanisms[J]. Annual Review of Phytopathology, 2015, 53: 445-470. doi: 10.1146/annurev-phyto-080614-115928 pmid: 26047563
[26]	Wiesner-Hanks T, Nelson R. Multiple disease resistance in plants[J]. Annual Review of Phytopathology, 2016, 54: 229-252. doi: 10.1146/annurev-phyto-080615-100037 pmid: 27296142
[27]	Mcintosh R A. Catalogue of gene symbols for wheat:2024[DB/OL]. https://graingenes.org/GG3/content/october-2024-wheat-gene-catalogue-2024-released-covering-all-wgc-curations.
[28]	Boden S A, Mcintosh R A, Uauy C, Krattinger S G, Dubcovsky J, Rogers W J, Xia X C, Badaeva E D, Bentley A R, Brown-Guedira G, Caccamo M, Cattivelli L, Chhuneja P, Cockram J, Contreras-Moreira B, Dreisigacker S, Edwards D, González F G, Guzmán C, Ikeda T M, et al. Updated guidelines for gene nomenclature in wheat[J]. Theoretical and Applied Genetics, 2023, 136(4): 72. doi: 10.1007/s00122-023-04253-w pmid: 36952017
[29]	Saintenac C, Cambon F, Aouini L, Verstappen E, Ghaffary S M T, Poucet T, Marande W, Berges H, Xu S, Jaouannet M, Favery B, Alassimone J, Sánchez-Vallet A, Faris J, Kema G, Robert O, Langin T. A wheat cysteine-rich receptor-like kinase confers broad-spectrum resistance against Septoria tritici blotch[J]. Nature Communications, 2021, 12: 433. doi: 10.1038/s41467-020-20685-0 pmid: 33469010
[30]	Hafeez A N, Chartrain L, Feng C, Cambon F, Clarke M, Griffiths S, Hayta S, Jiang M, Keller B, Kirby R, Kolodziej M C, Powell O R, Smedley M A, Steuernagel B, Xian W F, Wingen L U, Cheng S F, Saintenac C, Wulff B B, Brown J K. Septoria tritici blotch resistance gene Stb15 encodes a lectin receptor-like kinase[J]. Nature Plants, 2025, 11(3): 410-420. doi: 10.1038/s41477-025-01920-2 pmid: 40087541
[31]	Fu D, Uauy C, Distelfeld A, Blechl A, Epstein L, Chen X M, Fahima T, Dubcovsky J. A kinase-START gene confers temperature-dependent resistance to wheat stripe rust[J]. Science, 2009, 323(5919): 1357-1360. doi: 10.1126/science.1166289 pmid: 19228999
[32]	Klymiuk V, Yaniv E, Huang L, Raats D, Fatiukha A, Chen S S, Feng L H, Frenkel Z, Krugman T, Lidzbarsky G, Chang W, Jääskeläinen M J, Schudoma C, Paulin L, Laine P, Bariana H, Sela H, Saleem K, Sørensen C K, Hovmøller M S, et al. Cloning of the wheat Yr15 resistance gene sheds light on the plant tandem kinase-pseudokinase family[J]. Nature Communications, 2018, 9: 3735. doi: 10.1038/s41467-018-06138-9
[33]	Lu P, Guo L, Wang Z, Li B B, Li J, Li Y H, Qiu D, Shi W Q, Yang L J, Wang N, Guo G H, Xie J Z, Wu Q H, Chen Y X, Li M M, Zhang H Z, Dong L L, Zhang P P, Zhu K Y, Yu D Z, et al. A rare gain of function mutation in a wheat tandem kinase confers resistance to powdery mildew[J]. Nature Communications, 2020, 11: 680. doi: 10.1038/s41467-020-14294-0 pmid: 32015344
[34]	Wang Y, Abrouk M, Gourdoupis S, Koo D H, Karafiátová M, Molnár I, Holušová K, Doležel J, Athiyannan N, Cavalet-Giorsa E, Jaremko Ł, Poland J, Krattinger S G. An unusual tandem kinase fusion protein confers leaf rust resistance in wheat[J]. Nature Genetics, 2023, 55(6): 914-920. doi: 10.1038/s41588-023-01401-2
[35]	Krattinger S G, Lagudah E S, Spielmeyer W, Singh R P, Huerta- Espino J, McFadden H, Bossolini E, Selter L L, Keller B. A putative ABC transporter confers durable resistance to multiple fungal pathogens in wheat[J]. Science, 2009, 323(5919): 1360-1363. doi: 10.1126/science.1166453 pmid: 19229000
[36]	Moore J W, Herrera-Foessel S, Lan C, Schnippenkoetter W, Ayliffe M, Huerta-Espino J, Lillemo M, Viccars L, Milne R, Periyannan S, Kong X Y, Spielmeyer W, Talbot M, Bariana H, Patrick J W, Dodds P, Singh R, Lagudah E. A recently evolved hexose transporter variant confers resistance to multiple pathogens in wheat[J]. Nature Genetics, 2015, 47(12): 1494-1498. doi: 10.1038/ng.3439 pmid: 26551671
[37]	Jones J D G, Staskawicz B J, Dangl J L. The plant immune system: From discovery to deployment[J]. Cell, 2024, 187(9): 2095-2116. doi: 10.1016/j.cell.2024.03.045 pmid: 38670067
[38]	Yuan M, Ngou B P, Ding P, Xin X F. PTI-ETI crosstalk: An integrative view of plant immunity[J]. Current Opinion in Plant Biology, 2021, 62: 102030. doi: 10.1016/j.pbi.2021.102030
[39]	Förderer A, Li E, Lawson A W, Deng Y, Sun Y, Logemann E, Zhang X, Wen J, Han Z, Chang J, Chen Y, Schulze-Lefert P, Chai J. A wheat resistosome defines common principles of immune receptor channels[J]. Nature, 2022, 610(7932): 532-539. doi: 10.1038/s41586-022-05231-w
[40]	Guo G, Zhao H, Bai K, Lu J, Wu Q, Lu L, Zhang Y, Dong L, Li G, Chen Y, Hou Y, Lu P, Li M, Zhang H, Wang G, Zhu K, Huang B, Cui X, Fu H, Hu C. An activated wheat CCG10-NLR immune receptor forms an octameric resistosome[J]. Cell, 2026, 189: 2955-2970. doi: 10.1016/j.cell.2026.02.024
[41]	Faris J D, Zhang Z, Lu H, Lu S, Reddy L, Cloutier S, Fellers J P, Meinhardt S W, Rasmussen J B, Xu S, Oliver R P, Simons K J, Friesen T L. A unique wheat disease resistance-like gene governs effector-triggered susceptibility to necrotrophic pathogens[J]. Proceedings of the National Academy of Sciences of the United States of America, 2010, 107(30): 13544-13549. doi: 10.1073/pnas.1004090107 pmid: 20624958
[42]	Hurni S, Brunner S, Stirnweis D, Herren G, Peditto D, McIntosh R A, Keller B. The powdery mildew resistance gene Pm8 derived from rye is suppressed by its wheat ortholog Pm3[J]. The Plant Journal, 2014, 79(6): 904-913. doi: 10.1111/tpj.2014.79.issue-6
[43]	Singh S P, Hurni S, Ruinelli M, Brunner S, Sanchez-Martin J, Krukowski P, Peditto D, Buchmann G, Zbinden H, Keller B. Evolutionary divergence of the rye Pm17 and Pm8 resistance genes reveals ancient diversity[J]. Plant Molecular Biology, 2018, 98(3): 249-260. doi: 10.1007/s11103-018-0780-3
[44]	Yan X, Li M, Zhang P, Yin G, Zhang H, Gebrewahid T W, Zhang J, Dong L, Liu D, Liu Z, Li Z. High-temperature wheat leaf rust resistance gene Lr13 exhibits pleiotropic effects on hybrid necrosis[J]. Molecular Plant, 2021, 14(7): 1029-1032. doi: 10.1016/j.molp.2021.05.009
[45]	Hewitt T, Zhang J, Huang L, Upadhyaya N, Li J, Park R, Hoxha S, McIntosh R, Lagudah E, Zhang P. Wheat leaf rust resistance gene Lr13 is a specific Ne2 allele for hybrid necrosis[J]. Molecular Plant, 2021, 14(7): 1025-1028. doi: 10.1016/j.molp.2021.05.010
[46]	Athiyannan N, Abrouk M, Boshoff W H P, Cauet S, Rodde N, Kudrna D, Mohammed N, Bettgenhaeuser J, Botha K S, Derman S S, Wing R A, Prins R, Krattinger S G. Long-read genome sequencing of bread wheat facilitates disease resistance gene cloning[J]. Nature Genetics, 2022, 54(3): 227-231. doi: 10.1038/s41588-022-01022-1 pmid: 35288708
[47]	Xie J, Guo G, Wang Y, Hu T, Wang L, Li J, Qiu D, Li Y, Wu Q, Lu P, Chen Y, Dong L, Li M, Zhang H, Zhang P, Zhu K, Li B, Deal K R, Huo N, Zhang Y, et al. A rare single nucleotide variant in Pm5e confers powdery mildew resistance in common wheat[J]. New Phytologist, 2020, 228(3): 1011-1026. doi: 10.1111/nph.v228.3
[48]	Sun W, Dong H, Liu S, Ma S, Zeng Q, Li J, Ke K, Yue W, Zhang W, Fang X, Han J, Zhou X, Zhao J, Guo G, Li G, Cao X, Zheng W, Li C, Kang Z, Han D. The wheat Yr6 locus, allelic to Pm5, harbors an NLR gene pair conferring stripe rust resistance[J]. Plant Communications, 2026, 7(1): 101541. doi: 10.1016/j.xplc.2025.101541
[49]	Huang S, Zhang L, Liang J, Ouyang Y, Yan Y, Ju H, Wang Y, Zhang H, Liu T, Tang C, Wang X, Wang Y. Genome-based mutant RNA mapping identifies an NLR pair underlying Yr6-mediated stripe rust resistance in wheat[J]. Plant Communications, 2026, 7(1): 101542. doi: 10.1016/j.xplc.2025.101542
[50]	Steuernagel B, Periyannan S K, Hernández-Pinzón I, Witek K, Rouse M N, Yu G T, Hatta A, Ayliffe M, Bariana H, Jones J D, Lagudah E S, Wulff B B. Rapid cloning of disease-resistance genes in plants using mutagenesis and sequence capture[J]. Nature Biotechnology, 2016, 34(6): 652-655. doi: 10.1038/nbt.3543 pmid: 27111722
[51]	Jin Y, Li W, Li Y, Li D, Yan H, Chen S, Han G, Xiao B, Li B, Lu P, Chen Y, Wang Y, Su F, Yu N, Zhang J, Wang K, Liu Z, He H, Liu C, Wu Q, et al. Pm37 as a susceptible Sr22 allele confers resistance to wheat powdery mildew and leaf rust[J]. Nature Communications, 2026, 17: 3165. doi: 10.1038/s41467-026-69717-1
[52]	Sharma D, Avni R, Gutierrez-Gonzalez J, Kumar R, Sela H, Prusty M R, Shatil-Cohen A, Molnár I, Holušová K, Said M, Doležel J, Millet E, Khazan-Kost S, Landau U, Bethke G, Sharon O, Ezrati S, Ronen M, Maatuk O, Eilam T, et al. A single NLR gene confers resistance to leaf and stripe rust in wheat[J]. Nature Communications, 2024, 15: 9925. doi: 10.1038/s41467-024-54068-6
[53]	Wulff B B, Liu Z. Good things come in pairs: Crop disease resistance from sensor-helper to sensor-executor pairs[J]. The Crop Journal, 2025, 13(6): 1655-1659. doi: 10.1016/j.cj.2025.10.003
[54]	Zhang H, Li M, Wang G, Zhu K, Guo G, Fu H, Hu C, Chu Z, Hu J, Wu Q, Chen Y, Qiu D, Xie J, Li D, Li B, Li W, Dong L, Hou Y, Cui X, Huang B, et al. Paired NLRs originated from Triticum dicoccoides coordinately confer resistance to powdery mildew in wheat[J]. Nature Communications, 2025, 16: 9040. doi: 10.1038/s41467-025-64049-y
[55]	Yang Z, Liu N, Xie X, Wei W, Bai Y, Sun J, Pan W, Yang J, Wang W, Xie X, Saqlain M, Kang H, Li B, Hu Z, Gou J, Guo W, Song S, Ma J, Fahima T, Sun Q, et al. Two complementary NLRs from wild emmer wheat confer powdery mildew resistance[J]. Nature Communications, 2025, 16: 9041. doi: 10.1038/s41467-025-64052-3
[56]	He H, Tang Q, Zhang Q, Zhu S, Lv S, Bao Y, Liang J, Wang J, Wang J, Xu H, Cavalet-Giorsa E, Krattinger S G, Li H, Wu C, Gao A, Wang Y. An NLR pair in the Pm68 locus confers powdery mildew resistance in durum and common wheat[J]. Nature Communications, 2025, 16: 9039. doi: 10.1038/s41467-025-64048-z
[57]	Klymiuk V, Wiebe K, Chawla H S, Ens J, Subramaniam R, Pozniak C J. Coordinated function of paired NLRs confers Yr84-mediated stripe rust resistance in wheat[J]. Nature Genetics, 2025, 57(6): 1535-1542. doi: 10.1038/s41588-025-02203-4
[58]	Hu Y, Li M, Li Y, Du L, Xie R, Ni F, Xia C, Wang K, Huang Y, Xu B, Li Y, Jiang Y, Hao M, Jiang B, Ning S, Yuan Z, Feng L, Zhang L, Chen S, Wu B, et al. A head-to-head NLR gene pair from wild emmer confers stripe rust resistance in wheat[J]. Nature Genetics, 2025, 57(6): 1543-1552. doi: 10.1038/s41588-025-02207-0
[59]	Zhu K, Li M, Dong L, Zhang H, Zhang D, Lu P, Wu Q, Xie J, Chen Y, Guo G, Zhang P, Li B, Li W, Dong L, Hou Y, Yang Y, Qiu D, Wang G, Huang B, Cui X, et al. An atypical NLR pair TdCNL1/TdCNL5 from wild emmer confers powdery mildew resistance in wheat[J]. Nature Genetics, 2025, 57(6): 1553-1562. doi: 10.1038/s41588-025-02208-z
[60]	Guo G, Bai K, Hou Y, Gong Z, Zhang H, Wu Q, Lu P, Li M, Dong L, Xie J, Chen Y, Zhang P, Zhu K, Li B, Li W, Dong L, Yang Y, Qiu D, Wang G, Ahn H K, et al. The wheat NLR pair RXL/Pm5e confers resistance to powdery mildew[J]. Plant Biotechnology Journal, 2025, 23(4): 1260-1276. doi: 10.1111/pbi.v23.4
[61]	Marchal C, Zhang J, Zhang P, Fenwick P, Steuernagel B, Adamski N M, Boyd L, McIntosh R, Wulff B B, Berry S, Lagudah E, Uauy C. BED-domain-containing immune receptors confer diverse resistance spectra to yellow rust[J]. Nature Plants, 2018, 4(9): 662-668. doi: 10.1038/s41477-018-0236-4 pmid: 30150615
[62]	Wu J, Ma S, Niu J, Sun W, Dong H, Zheng S, Zhao J, Liu S, Yu R, Li Y, Han J, Wang Y, Chen T, Zhang C, Zhang W, Ding B, Chang L, Xue W, Zheng W, Li C, et al. Genomics-driven discovery of superior alleles and genes for yellow rust resistance in wheat[J]. Nature Genetics, 2025, 57(8): 2017-2027. doi: 10.1038/s41588-025-02259-2
[63]	Wang H, Zou S, Li Y, Lin F, Tang D. An ankyrin-repeat and WRKY-domain-containing immune receptor confers stripe rust resistance in wheat[J]. Nature Communications, 2020, 11: 1353. doi: 10.1038/s41467-020-15139-6 pmid: 32170056
[64]	Kema G H, Gohari A M, Aouini L, Gibriel H A, Ware S B, van den Bosch F, Manning-Smith R, Alonso-Chavez V, Helps J, Ben M’Barek S, Mehrabi R, Diaz-Trujillo C, Zamani E, Schouten H J, van der Lee T A, Waalwijk C, de Waard M A, de Wit P J, Verstappen E C, Thomma B P, et al. Stress and sexual reproduction affect the dynamics of the wheat pathogen effector AvrStb6 and strobilurin resistance[J]. Nature Genetics, 2018, 50(3): 375-380. doi: 10.1038/s41588-018-0052-9 pmid: 29434356
[65]	Shi G, Zhang Z, Friesen T L, Raats D, Fahima T, Brueggeman R S, Lu S, Trick H N, Liu Z, Chao W, Frenkel Z, Xu S, Rasmussen J B, Faris J D. The hijacking of a receptor kinase-driven pathway by a wheat fungal pathogen leads to disease[J]. Science Advances, 2016, 2(10): e1600822. doi: 10.1126/sciadv.1600822
[66]	Fan A, Wei L, Zhang X, Liu J, Sun L, Xiao J, Wang Y, Wang H, Hua J, Singh R P, Wang Z, Wang X. Heterologous expression of the Haynaldia villosa pattern-recognition receptor CERK1-V in wheat increases resistance to three fungal diseases[J]. The Crop Journal, 2022, 10(6): 1733-1745. doi: 10.1016/j.cj.2022.02.005
[67]	Powell O R, Guzmán-Vega F J, Yu D, Wang Y, Lu P, Arold S T, Liu Z, Banfield M J, Wulff B B H, Chen R. The emerging role of kinase fusion proteins in cereal immunity[J]. Nature Genetics, 2026, 58(4): 695-703. doi: 10.1038/s41588-026-02515-z
[68]	Reveguk T, Fatiukha A, Potapenko E, Reveguk I, Sela H, Klymiuk V, Li Y, Pozniak C, Wicker T, Coaker G, Fahima T. Tandem kinase proteins across the plant kingdom[J]. Nature Genetics, 2025, 57(1): 254-262. doi: 10.1038/s41588-024-02032-x pmid: 39779952
[69]	Sung Y, Li Y, Bernasconi Z, Baik S, Asuke S, Keller B, Fahima T, Coaker G. Wheat tandem kinase RWT4 directly binds a fungal effector to activate defense[J]. Nature Genetics, 2025, 57(5): 1238-1249. doi: 10.1038/s41588-025-02162-w
[70]	Lu P, Zhang G, Li J, Gong Z, Wang G, Dong L, Zhang H, Guo G, Su M, Wang K, Wang Y, Zhu K, Wu Q, Chen Y, Li M, Huang B, Li B, Li W, Dong L, Hou Y, et al. A wheat tandem kinase and NLR pair confers resistance to multiple fungal pathogens[J]. Science, 2025, 387(6741): 1418-1424. doi: 10.1126/science.adp5469 pmid: 40146830
[71]	Chen R, Chen J, Powell O R, Outram M A, Arndell T, Gajendiran K, Wang Y, Lubega J, Xu Y, Ayliffe M A, Blundell C, Figueroa M, Sperschneider J, Vanhercke T, Kanyuka K, Tang D, Zhong G, Gardener C, Yu G, Gourdoupis S, et al. A wheat tandem kinase activates an NLR to trigger immunity[J]. Science, 2025, 387(6741): 1402-1408. doi: 10.1126/science.adp5034 pmid: 40146821
[72]	Dracatos P M, Lu J, Sánchez-Martín J, Wulff B B. Resistance that stacks up: Engineering rust and mildew disease control in the cereal crops wheat and barley[J]. Plant Biotechnology Journal, 2023, 21(10): 1938-1951. doi: 10.1111/pbi.14106 pmid: 37494504
[73]	He H, Chen Z, Fan R, Zhang J, Zhu S, Wang J, Zhang Q, Gao A, Gong S, Zhang L, Li Y, Zhao Y, Krattinger S G, Shen Q, Li H, Wang Y. A kinase fusion protein from Aegilops longissima confers resistance to wheat powdery mildew[J]. Nature Communications, 2024, 15: 6512. doi: 10.1038/s41467-024-50909-6
[74]	Li H, Men W, Ma C, Liu Q, Dong Z, Tian X, Wang C, Liu C, Gill H S, Ma P, Zhang Z, Liu B, Zhao Y, Sehgal S K, Liu W. Wheat powdery mildew resistance gene Pm13 encodes a mixed lineage kinase domain-like protein[J]. Nature Communications, 2024, 15: 2449. doi: 10.1038/s41467-024-46814-7
[75]	Sánchez-Martín J, Widrig V, Herren G, Wicker T, Zbinden H, Gronnier J, Spörri L, Praz C R, Heuberger M, Kolodziej M C, Isaksson J, Steuernagel B, Karafiátová M, Doležel J, Zipfel C, Keller B. Wheat Pm4 resistance to powdery mildew is controlled by alternative splice variants encoding chimeric proteins[J]. Nature Plants, 2021, 7(3): 327-341. doi: 10.1038/s41477-021-00869-2 pmid: 33707738
[76]	Bernasconi Z, Herger A G, Caro M D, Kunz L, Müller M C, Stirnemann U, Outram M A, Widrig V, Neidhart M, Isaksson J, Schudel S, Rösli S, Wicker T, Bender K W, Zipfel C, Dodds P N, Figueroa M, Sánchez-Martín J, Keller B. Virulence on Pm4 kinase-based resistance is determined by two divergent wheat powdery mildew effectors[J]. Nature Plants, 2026, 12(1): 164-178. doi: 10.1038/s41477-025-02180-w
[77]	Gou J, Li K, Wu K, Wang X, Lin H, Cantu D, Uauy C, Dobon-Alonso A, Midorikawa T, Inoue K, Sánchez J, Fu D, Blechl A, Wallington E, Fahima T, Meeta M, Epstein L, Dubcovsky J. Wheat stripe rust resistance protein WKS1 reduces the ability of the thylakoid- associated ascorbate peroxidase to detoxify reactive oxygen species[J]. The Plant Cell, 2015, 27(6): 1755-1770. doi: 10.1105/tpc.114.134296
[78]	Wang S, Li Q, Wang J, Yan Y, Zhang G, Yan Y, Zhang H, Wu J, Chen F, Wang X, Kang Z, Dubcovsky J, Gou J. YR36/WKS1- mediated phosphorylation of Psbo, an extrinsic member of photosystem II, inhibits photosynthesis and confers stripe rust resistance in wheat[J]. Molecular Plant, 2019, 12(12): 1639-1650. doi: 10.1016/j.molp.2019.10.005
[79]	Arora S, Steed A, Goddard R, Gaurav K, O’Hara T, Schoen A, Rawat N, Elkot A F, Korolev A V, Chinoy C, Nicholson M H, Asuke S, Antoniou-Kourounioti R, Steuernagel B, Yu G, Awal R, Forner- Martínez M, Wingen L, Baggs E, Clarke J, et al. A wheat kinase and immune receptor form host-specificity barriers against the blast fungus[J]. Nature Plants, 2023, 9(3): 385-392. doi: 10.1038/s41477-023-01357-5 pmid: 36797350
[80]	O’Hara T, Steed A, Goddard R, Gaurav K, Arora S, Quiroz-Chávez J, Ramírez-González R, Badgami R, Gilbert D, Sánchez-Martín J, Wingen L, Feng C, Jiang M, Cheng S F, Dreisigacker S, Keller B, Wulff B B H, Uauy C, Nicholson P. The wheat powdery mildew resistance gene Pm4 also confers resistance to wheat blast[J]. Nature Plants, 2024, 10(6): 984-993. doi: 10.1038/s41477-024-01718-8
[81]	Asuke S, Morita K, Shimizu M, Abe F, Terauchi R, Nago C, Takahashi Y, Shibata M, Yoshioka M, Iwakawa M, Kishi-Kaboshi M, Su Z, Nasuda S, Handa H, Fujita M, Tougou M, Hatta K, Mori N, Matsuoka Y, Kato K, et al. Evolution of wheat blast resistance gene Rmg8 accompanied by differentiation of variants recognizing the powdery mildew fungus[J]. Nature Plants, 2024, 10(6): 971-983. doi: 10.1038/s41477-024-01711-1
[82]	Zhao Y, Dong Z, Miao J, Liu Q, Ma C, Tian X, He J, Bi H, Yao W, Li T, Gill H S, Zhang Z, Cao A, Liu B, Li H, Sehgal S K, Liu W. Pm57 from Aegilops searsii encodes a tandem kinase protein and confers wheat powdery mildew resistance[J]. Nature Communications, 2024, 15: 4796. doi: 10.1038/s41467-024-49257-2 pmid: 38839783
[83]	Deppe J P, Rabbat R, Hörtensteiner S, Keller B, Martinoia E, Lopéz-Marqués R L. The wheat ABC transporter Lr34 modifies the lipid environment at the plasma membrane[J]. Journal of Biological Chemistry, 2018, 293(48): 18667-18679. doi: 10.1074/jbc.RA118.002532 pmid: 30327425
[84]	Krattinger S G, Kang J, Bräunlich S, Boni R, Chauhan H, Selter L L, Robinson M D, Schmid M W, Wiederhold E, Hensel G, Kumlehn J, Sucher J, Martinoia E, Keller B. Abscisic acid is a substrate of the ABC transporter encoded by the durable wheat disease resistance gene Lr34[J]. New Phytologist, 2019, 223(2): 853-866. doi: 10.1111/nph.15815 pmid: 30913300
[85]	Milne R J, Dibley K E, Bose J, Ashton A R, Ryan P R, Tyerman S D, Lagudah E S. Expression of the wheat multipathogen resistance hexose transporter Lr67res is associated with anion fluxes[J]. Plant Physiology, 2023, 192(2): 1254-1267. doi: 10.1093/plphys/kiad104 pmid: 36806945
[86]	Milne R J, Dibley K E, Bose J, Riaz A, Zhang J P, Schnippenkoetter W, Ashton A R, Ryan P R, Tyerman S D, Lagudah E S. Dissecting the causal polymorphism of the Lr67res multipathogen resistance gene[J]. Journal of Experimental Botany, 2024, 75(13): 3877-3890. doi: 10.1093/jxb/erae164
[87]	Kolodziej M C, Singla J, Sánchez-Martín J, Zbinden H, Šimková H, Karafiátová M, Doležel J, Gronnier J, Poretti M, Glauser G, Zhu W S, Köster P, Zipfel C, Wicker T, Krattinger S G, Keller B. A membrane-bound ankyrin repeat protein confers race-specific leaf rust disease resistance in wheat[J]. Nature Communications, 2021, 12: 956. doi: 10.1038/s41467-020-20777-x pmid: 33574268
[88]	Yue L, Wang L, Neuhäuser B, Zhang S, Herren G, Jung E, Kim G, Goto Y, Heuberger M, Ludewig U, Zipfel C, Keller B. Cytoplasmic calcium influx mediated by Lr14a regulates stomatal immunity against leaf rust in wheat[J]. Current Biology, 2025, 35(23): 5750-5761. doi: 10.1016/j.cub.2025.10.033
[89]	Wang H, Sun S, Ge W, Zhao L, Hou B, Wang K, Lyu Z, Chen L, Xu S, Guo J, Li M, Su P, Li X, Wang G, Bo C, Fang X, Zhuang W, Cheng X, Wu J, Dong L, et al. Horizontal gene transfer of Fhb7 from fungus underlies Fusarium head blight resistance in wheat[J]. Science, 2020, 368(6493): eaba5435.
[90]	He Y, Yang X, Xia X, Wang Y, Dong Y, Wu L, Jiang P, Zhang X, Jiang C, Ma H, Ma W, Liu C, Whitford R, Tucker M R, Zhang Z, Li G. A phase-separated protein hub modulates resistance to Fusarium head blight in wheat[J]. Cell Host & Microbe, 2024, 32(5): 710-726.
[91]	Kourelis J, van der Hoorn R A. Defended to the nines: 25 years of resistance gene cloning identifies nine mechanisms for R protein function[J]. The Plant Cell, 2018, 30(2): 285-299. doi: 10.1105/tpc.17.00579 pmid: 29382771
[92]	Eckardt N A. Plant disease susceptibility genes?[J]. The Plant Cell, 2002, 14(9): 1983-1986. doi: 10.1105/tpc.140910
[93]	Büschges R, Hollricher K, Panstruga R, Simons G, Wolter M, Frijters A, van Daelen R, van der Lee T, Diergaarde P, Groenendijk J, Töpsch S, Vos P, Salamini F, Schulze-Lefert P. The barley Mlo gene: A novel control element of plant pathogen resistance[J]. Cell, 1997, 88(5): 695-705. doi: 10.1016/s0092-8674(00)81912-1 pmid: 9054509
[94]	Kim M C, Panstruga R, Elliott C, Müller J, Devoto A, Yoon H W, Park H C, Cho M J, Schulze-Lefert P. Calmodulin interacts with MLO protein to regulate defence against mildew in barley[J]. Nature, 2002, 416(6879): 447-451. doi: 10.1038/416447a
[95]	Piffanelli P, Zhou F, Casais C, Orme J, Jarosch B, Schaffrath U, Collins N C, Panstruga R, Schulze-Lefert P. The barley MLO modulator of defense and cell death is responsive to biotic and abiotic stress stimuli[J]. Plant Physiology, 2002, 129(3): 1076-1085. doi: 10.1104/pp.010954 pmid: 12114562
[96]	Wang Y, Cheng X, Shan Q, Zhang Y, Liu J, Gao C, Qiu J. Simultaneous editing of three homoeoalleles in hexaploid bread wheat confers heritable resistance to powdery mildew[J]. Nature Biotechnology, 2014, 32(9): 947-951. doi: 10.1038/nbt.2969 pmid: 25038773
[97]	Li S, Lin D, Zhang Y, Deng M, Chen Y, Lv B, Li B, Lei Y, Wang Y, Zhao L, Liang Y, Liu J, Chen K, Liu Z, Xiao J, Qiu J, Gao C. Genome-edited powdery mildew resistance in wheat without growth penalties[J]. Nature, 2022, 602(7897): 455-460. doi: 10.1038/s41586-022-04395-9
[98]	Wang N, Tang C, Fan X, He M, Gan P, Zhang S, Hu Z, Wang X, Yan T, Shu W, Yu L, Zhao J, He J, Li L, Wang J, Huang X, Huang L, Zhou J, Kang Z, Wang X. Inactivation of a wheat protein kinase gene confers broad-spectrum resistance to rust fungi[J]. Cell, 2022, 185(16): 2961-2974. doi: 10.1016/j.cell.2022.06.027
[99]	Farooq M, Frei M, Zeibig F, Pantha S, Özkan H, Kilian B, Siddique K H. Back into the wild: Harnessing the power of wheat wild relatives for future crop and food security[J]. Journal of Experimental Botany, 2026, 77(9): 2645-2665. doi: 10.1093/jxb/eraf141
[100]	刘成, 韩冉, 汪晓璐, 宫文萍, 程敦公, 曹新有, 刘爱峰, 李豪圣, 刘建军. 小麦远缘杂交现状、抗病基因转移及利用研究进展[J]. 中国农业科学, 2020, 53(7): 1287-1308. DOI: 10.3864/j.issn.0578-1752.2020.07.001.
	Liu C, Han R, Wang X L, Gong W P, Cheng D G, Cao X Y, Liu A F, Li H S, Liu J J. Research progress of wheat wild hybridization, disease resistance genes transfer and utilization[J]. Scientia Agricultura Sinica, 2020, 53(7): 1287-1308. DOI: 10.3864/j.issn.0578-1752.2020.07.001. (in Chinese)
[101]	Arora S, Steuernagel B, Gaurav K, Chandramohan S, Long Y, Matny O, Johnson R, Enk J, Periyannan S, Singh N, Hatta M, Athiyannan N, Cheema J, Yu G, Kangara N, Ghosh S, Szabo L J, Poland J, Bariana H, Jones J D, et al. Resistance gene cloning from a wild crop relative by sequence capture and association genetics[J]. Nature Biotechnology, 2019, 37(2): 139-143. doi: 10.1038/s41587-018-0007-9 pmid: 30718880
[102]	Cavalet-Giorsa E, González-Muñoz A, Athiyannan N, Holden S, Salhi A, Gardener C, Quiroz-Chávez J, Rustamova S M, Elkot A F, Patpour M, Rasheed A, Mao L, Lagudah E S, Periyannan S K, Sharon A, Himmelbach A, Reif J C, Knauft M, Mascher M, Stein N, et al. Origin and evolution of the bread wheat D genome[J]. Nature, 2024, 633(8031): 848-855. doi: 10.1038/s41586-024-07808-z
[103]	Brabham H J, Hernández-Pinzón I, Yanagihara C, Ishikawa N, Komori T, Matny O N, Hubbard A, Witek K, Feist A, Numazawa H, Green P, Dreiseitl A, Takemori N, Komari T, Freedman R P, Steffenson B, van Esse H P, Moscou M J. Discovery of functional NLRs using expression level, high-throughput transformation and large-scale phenotyping[J]. Nature Plants, 2025, 11(10): 2100-2114. doi: 10.1038/s41477-025-02110-w
[104]	Yang J, Li H, Li M, Song R, Shen T, Wang G, Xu D, Hao M, Jia A, Rehman S U, Hua L, Liang Y, Chi C, Lan C, Deng X, Dubcovsky J, Song B, Wang X, Chen S. Genome-assisted identification of wheat leaf rust resistance gene Lr.ace-4A/Lr30[J]. Nature Communications, 2025, 16: 9339. doi: 10.1038/s41467-025-64428-5
[105]	Zhang W, Chen S, Abate Z, Nirmala J, Rouse M N, Dubcovsky J. Identification and characterization of Sr13, a tetraploid wheat gene that confers resistance to the Ug99 stem rust race group[J]. Proceedings of the National Academy of Sciences of the United States of America, 2017, 114(45): E9483-E9492.
[106]	Li M, Dong L, Zhu K, Wu Q, Chen Y, Lu P, Guo G, Zhang H, Zhang P, Li B, Li W, Yang Y, Hou Y, Cui X, Li H, Dong L, Zhao Y, Liu Z. Provoking a silent R gene in wheat genome confers resistance to powdery mildew[J]. Plant Biotechnology Journal, 2022, 20(11): 2039-2041. doi: 10.1111/pbi.13903 pmid: 35904481
[107]	Tamborski J, Seong K, Liu F, Staskawicz B J, Krasileva K V. Altering specificity and autoactivity of plant immune receptors Sr33 and Sr50 via a rational engineering approach[J]. Molecular Plant-Microbe Interactions, 2023, 36(7): 434-446. doi: 10.1094/MPMI-07-22-0154-R pmid: 36867580
[108]	Liu S, Xiang M, Wang X, Li J, Cheng X, Li H, Singh R P, Bhavani S, Huang S, Zheng W, Li C, Yuan F, Wu J, Han D, Kang Z, Zeng Q. Development and application of the GenoBaits WheatSNP16K array to accelerate wheat genetic research and breeding[J]. Plant Communications, 2025, 6(1): 101138. doi: 10.1016/j.xplc.2024.101138
[109]	Deng P, Du X, Wang Y, Yang X, Cheng X, Huang C, Li T, Chen C, Zhao J, Wang C, Liu X, Tian Z, Ji W. GenoBaits^®WheatplusEE: A targeted capture sequencing panel for quick and accurate identification of wheat-Thinopyrum derivatives[J]. Theoretical and Applied Genetics, 2024, 137(2): 36. doi: 10.1007/s00122-023-04538-0
[110]	Xiang M, Liu S, Wang X, Zhang M, Yan W, Wu J, Wang Q, Li C, Zheng W, He Y, Ge Y, Wang C, Kang Z, Han D, Zeng Q. Development of breeder chip for gene detection and molecular- assisted selection by target sequencing in wheat[J]. Molecular Breeding, 2023, 43(2): 13. doi: 10.1007/s11032-023-01359-3
[111]	Zhao J, Dong H, Han J, Ou J, Chen T, Wang Y, Liu S, Yu R, Zheng W, Li C, Kang Z, Han D, Zeng Q, Wang X, Ma S, Wu J. LWRR: Landscape of wheat rust resistance towards practical breeding design[J]. Stress Biology, 2025, 5(1): 25. doi: 10.1007/s44154-025-00232-x
[112]	孟祥宇, 刁邓超, 刘雅睿, 李云丽, 孙玉晨, 吴玮, 赵雯, 汪妤, 吴建辉, 李春莲, 曾庆东, 韩德俊, 郑炜君. 小麦新品种西农877高产稳产的遗传特性解析[J]. 作物学报, 2025, 51(5): 1261-1276. doi: 10.3724/SP.J.1006.2025.41064
	Meng X Y, Diao D C, Liu Y R, Li Y L, Sun Y C, Wu W, Zhao W, Wang Y, Wu J H, Li C L, Zeng Q D, Han D J, Zheng W J. Genetic analysis of high yield and yield stability characteristics of new wheat variety Xinong 877[J]. Acta Agronomica Sinica, 2025, 51(5): 1261-1276. (in Chinese) doi: 10.3724/SP.J.1006.2025.41064
[113]	李笑笑, 相明杰, 刘胜杰, 王晓婷, 李杰, 郑炜君, 吴建辉, 曾庆东, 康振生, 张传量, 王长发, 韩德俊. 结合0.1K前景和16K背景芯片快速解析小麦新品系的基因组结构和重要性状遗传基础[J]. 麦类作物学报, 2025, 45(1): 52-62.
	Li X X, Xiang M J, Liu S J, Wang X T, Li J, Zheng W J, Wu J H, Zeng Q D, Kang Z S, Zhang C L, Wang C F, Han D J. 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)
[114]	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. doi: 10.1016/j.xplc.2023.100549