Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (16): 3556-3576.doi: 10.3864/j.issn.0578-1752.2026.16.007

• PLANT PROTECTION • Previous Articles     Next Articles

Advances in Mechanisms and Innovative Improvement Strategies for Wheat Disease Resistance

WU JianHui1,3(), ZENG QingDong2,3, LIU ShengJie2,3, WANG XiaoJie2,3, HAN DeJun1,3, KANG ZhenSheng2,3   

  1. 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
  • Received:2026-03-22 Accepted:2026-06-06 Online:2026-08-16 Published:2026-08-17
  • Contact: WU JianHui

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

Fig. 1

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]"

Fig. 2

Types of wheat disease resistance"

Table 1

Cloned disease resistance genes in wheat"

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)

Fig. 3

Structural diversity and mediated immune defense mechanisms of wheat disease resistance genes"

Fig. 4

Future strategies for wheat disease resistance breeding"

[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
[1] WANG XiaoWei, DU FoLi, YAN HongCai, LANG ZhengDong, DANG ZhiJuan, LI BaoChun, WANG JunCheng, MA XiaoLe, WANG HuaJun, ZHANG Hong, YAO LiRong. Evaluation of Drought Resistance of 396 Spring Wheat Varieties at Grain Filling Stage and Maturity Stage [J]. Scientia Agricultura Sinica, 2026, 59(8): 1608-1621.
[2] WANG CaiYu, LIU XiaoLi, LI WenGuang, YANG WenPing, YANG ZhenPing, GAO ZhiQiang. Effects of Different Substitution Rates of Organic Fertilizers on Soil Multifunctionality and Its Microbial Driving Mechanisms [J]. Scientia Agricultura Sinica, 2026, 59(8): 1712-1726.
[3] ZHU Qi, JIA ZhenPeng, Tahir SHAH, XU ChenSheng, LI ZhiQi, LÜ HuiShuai, ZHU PengChao, WEI XiaoMin, HUANG DongLin, SUN YanNi, CAO WeiDong, GAO YaJun, WANG ZhaoHui, ZHANG DaBin. Green Manure Crops Combined with Enhanced-Efficiency Products Reduced Greenhouse Gas Emissions and Carbon Footprints in Dryland Wheat Fields [J]. Scientia Agricultura Sinica, 2026, 59(7): 1507-1522.
[4] LI WenHu, LI HaiFeng, DU YuPeng, DING YuLan, LUO YiNuo, LI YuKe, SHE WenTing, ZHANG Feng, TENG Yu, ZHANG SiQi, HUANG Cui, LI XiaoHan, LIU JinShan, WANG ZhaoHui. Regional Differences in Wheat Zinc Uptake and Translocation Responses to Soil Zinc Fertilization [J]. Scientia Agricultura Sinica, 2026, 59(5): 1034-1047.
[5] JIAO WenJuan, HE WanLong, GENG HongWei, BAI Bin, LI JianFeng, CHENG YuKun. Stripe Rust Resistance Evaluation and Molecular Characterization of Yr Genes for 155 Spring Wheat Varieties (Lines) [J]. Scientia Agricultura Sinica, 2026, 59(5): 937-950.
[6] CUI ShiYou, CHEN PengJun, MIAO YuanQing, HAN JiJun, SHEN JunMing. Development and Field Evaluation of Glyphosate-Resistant Wheat Germplasm Generated Through EMS Mutagenesis [J]. Scientia Agricultura Sinica, 2026, 59(4): 723-733.
[7] QIAN Jin, LI YingXue, WU Fang, ZOU XiaoChen. Improved Leaf Phosphorus Content Estimation of Winter Wheat Using Ensemble Hyperspectral Dimensionality Reduction Method [J]. Scientia Agricultura Sinica, 2026, 59(4): 781-792.
[8] KONG Yuan, CUI ShaSha, LI Mei, LI Jian, YANG SiYu, FANG Feng, LIU ShuaiShuai, LIU MingPing, ZENG Yan, GAO XingXiang, BAI LianYang. Spatiotemporal Distribution Dynamics of Five Grass Weed Species Including Lolium multiflorum in Winter Wheat Fields of the Huang- Huai-Hai Region [J]. Scientia Agricultura Sinica, 2026, 59(4): 807-823.
[9] WANG YongSheng, NIU Li, WANG ChangJie, MA LiHua, LIAN XiaoXiao, MENG YaXiong, MA XiaoLe, YAO LiRong, ZHANG Hong, YANG Ke, LI BaoChun, WANG HuaJun, SI ErJing, WANG JunCheng. Genome-Wide Association Study and Candidate Gene Identification for Thousand Grain Weight in Winter Wheat [J]. Scientia Agricultura Sinica, 2026, 59(3): 499-514.
[10] LI XinYi, LI JiaNing, YANG WenPing, XIA Qing, HUO YingRui, HAO ShiHang, HUANG TingMiao, REN YongKang, CHEN Jie, GAO ZhiQiang, YANG ZhenPing. Effects of Post-Anthesis Foliar Zinc Application on Zinc Nutrition in Colored-Grain Wheat [J]. Scientia Agricultura Sinica, 2026, 59(3): 515-527.
[11] XIAN QingLin, XIAO JianKe, GAO AQing, GAO LiChuang, LIU Yang. Effects of Planting Patterns Combined with Soil Moisture Measurement and Supplementary Irrigation on the Yield and Water Use Efficiency of Winter Wheat [J]. Scientia Agricultura Sinica, 2026, 59(3): 589-601.
[12] ZHANG ZhiYong, TAN ShiChao, XIONG ShuPing, MA XinMing, WEI YiHao, WANG XiaoChun. Effects of Annual Water and Nitrogen Optimization on Yield and Nitrogen Migration of Wheat-Maize Rotation System in Irrigation Area of Northern Henan [J]. Scientia Agricultura Sinica, 2026, 59(2): 336-353.
[13] ZHANG YanFei, WU ZiYue, XUE ZhiWei, MA DongYun, KANG GuoZhang, FENG Wei, XIE YingXin, JING RuiLian, WANG WanZhang, MAO XinGuo, WANG ChenYang. Functional Characterization and Molecular Marker Development of the Wheat Transcription Factor Gene TaWRKY65-3A [J]. Scientia Agricultura Sinica, 2026, 59(15): 3237-3251.
[14] WANG Qi, GUO ShuangQi, MEI LanXin, WU ChengDong, ZHENG ChangHong, ZHOU Qin, CAI Jian, WANG Xiao, Huang Mei, ZHONG YingXin, JIANG Dong. Regulation of Wheat Lodging Resistance Through Culm Structure Optimization and Synergistic by Uniconazole and Prohexadione Calcium [J]. Scientia Agricultura Sinica, 2026, 59(15): 3283-3301.
[15] MAO JiaQi, WU BangBang, ZHAO JiaJia, HAO YuQiong, ZHENG XingWei, WU JianHui, ZENG QingDong, LIU MinJie, ZHENG Jun. Evaluation of Rust Resistance in the Seedling Stage of Wheat Varieties and Disease-Resistant Gene Detection in Shanxi Province, China [J]. Scientia Agricultura Sinica, 2026, 59(14): 3006-3021.
Viewed
Full text


Abstract

Cited

  Shared   
  Discussed   
No Suggested Reading articles found!