Lehti-Shiu, M. D. & Shiu, S. H. Diversity, classification and function of the plant protein kinase superfamily. Philos. Trans. R. Soc. Lond. B Biol. Sci. 367, 2619–2639 (2012).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Jamieson, P. A., Shan, L. & He, P. Plant cell surface molecular cypher: receptor-like proteins and their roles in immunity and development. Plant Sci. 274, 242–251 (2018).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

DeFalco, T. A. & Zipfel, C. Molecular mechanisms of early plant pattern-triggered immune signaling. Mol. Cell 81, 3449–3467 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Yu, X., Feng, B., He, P. & Shan, L. From chaos to harmony: responses and signaling upon microbial pattern recognition. Annu. Rev. Phytopathol. 55, 109–137 (2017).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Kourelis, J. & van der Hoorn, R. A. L. Defended to the nines: 25 years of resistance gene cloning identifies nine mechanisms for R protein function. Plant Cell. 30, 285–299 (2018).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Liang, X. et al. A Phytophthora capsici RXLR effector targets and inhibits the central immune kinases to suppress plant immunity. New Phytol. 232, 264–278 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Wan, C. et al. A serine-rich effector from the stripe rust pathogen targets a Raf-like kinase to suppress host immunity. Plant Physiol. 190, 762–778 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Chen, R., Gajendiran, K. & Wulff, B. B. H. R we there yet? Advances in cloning resistance genes for engineering immunity in crop plants. Curr. Opin. Plant Biol. 77, 102489 (2024).

Article 
CAS 
PubMed 

Google Scholar
 

Ngou, B. P. M., Ding, P. & Jones, J. D. G. Thirty years of resistance: zig-zag through the plant immune system. Plant Cell 34, 1447–1478 (2022).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Ngou, B. P. M., Ahn, H. K., Ding, P. & Jones, J. D. G. Mutual potentiation of plant immunity by cell-surface and intracellular receptors. Nature 592, 110–115 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Förderer, A. et al. A wheat resistosome defines common principles of immune receptor channels. Nature 610, 532–539 (2022).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Bi, G. et al. The ZAR1 resistosome is a calcium-permeable channel triggering plant immune signaling. Cell 184, 3528–3541 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Dangl, J. L. & Jones, J. D. G. Plant pathogens and integrated defence responses to infection. Nature 411, 826–833 (2001).

Article 
CAS 
PubMed 

Google Scholar
 

Sheikh, A. H. et al. Dynamic changes of the Prf/Pto tomato resistance complex following effector recognition. Nat. Commun. 14, 2568 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Van der Hoorn, R. A. L. & Kamoun, S. From guard to decoy: a new model for perception of plant pathogen effectors. Plant Cell 20, 2009–2017 (2008).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Marchal, C., Michalopoulou, V. A., Zou, Z., Cevik, V. & Sarris, P. F. Show me your ID: NLR immune receptors with integrated domains in plants. Essays Biochem. 66, 527–539 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Guo, L. et al. Specific recognition of two MAX effectors by integrated HMA domains in plant immune receptors involves distinct binding surfaces. Proc. Natl Acad. Sci. USA 115, 11637–11642 (2018).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Wang, Y. et al. An optimized disease resistance gene cloning workflow for wheat. Nat. Commun. 16, 4904 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Faris, J. D. et al. A unique wheat disease resistance-like gene governs effector-triggered susceptibility to necrotrophic pathogens. Proc. Natl Acad. Sci. USA 107, 13544–13549 (2010).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Goh, F. J., Huang, C. Y., Derevnina, L. & Wu, C. H. NRC immune receptor networks show diversified hierarchical genetic architecture across plant lineages. Plant Cell 36, 3399–3418 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Liu, Y., Hou, S. & Chen, S. Kinase fusion proteins: intracellular R-proteins in plant immunity. Trends Plant Sci. 29, 278–282 (2024).

Article 
PubMed 

Google Scholar
 

Klymiuk, V. et al. Cloning of the wheat Yr15 resistance gene sheds light on the plant tandem kinase-pseudokinase family. Nat. Commun. 9, 3735 (2018).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Fahima, T. & Coaker, G. Pathogen perception and deception in plant immunity by kinase fusion proteins. Nat. Genet. 55, 908–909 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Powell, O. R., Gilbert, D. & Wulff, B. B. H. An extended β-finger motif is near-ubiquitous in kinase fusion proteins. Preprint at bioRxiv https://doi.org/10.1101/2025.07.14.664643 (2025).

Segovia, V. et al. Yr36 confers partial resistance at temperatures below 18 °C to U.K. isolates of Puccinia striiformis. Phytopathology 104, 871–878 (2014).

Article 
PubMed 

Google Scholar
 

Chen, R. et al. A wheat tandem kinase activates an NLR to trigger immunity. Science 387, 1402–1408 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Lu, P. et al. A wheat tandem kinase and NLR pair confers resistance to multiple fungal pathogens. Science 387, 1418–1424 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Zhang, Z. et al. A protein kinase-major sperm protein gene hijacked by a necrotrophic fungal pathogen triggers disease susceptibility in wheat. Plant J. 106, 720–732 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Sánchez-Martín, J. et al. Wheat Pm4 resistance to powdery mildew is controlled by alternative splice variants encoding chimeric proteins. Nat. Plants 7, 327–341 (2021).

Article 
PubMed 
PubMed Central 

Google Scholar
 

O’Hara, T. et al. The wheat powdery mildew resistance gene Pm4 also confers resistance to wheat blast. Nat. Plants 10, 984–993 (2024).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Asuke, S. et al. Evolution of wheat blast resistance gene Rmg8 accompanied by differentiation of variants recognizing the powdery mildew fungus. Nat. Plants 10, 971–983 (2024).

Article 
CAS 
PubMed 

Google Scholar
 

Arora, S. et al. A wheat kinase and immune receptor form host-specificity barriers against the blast fungus. Nat. Plants 9, 385–392 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Zhao, Y. et al. Pm57 from Aegilops searsii encodes a tandem kinase protein and confers wheat powdery mildew resistance. Nat. Commun. 15, 4796 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Lu, P. et al. A rare gain of function mutation in a wheat tandem kinase confers resistance to powdery mildew. Nat. Commun. 11, 680 (2020).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Reveguk, T. et al. Tandem kinase proteins across the plant kingdom. Nat. Genet. 57, 254–262 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Cesari, S. Multiple strategies for pathogen perception by plant immune receptors. New Phytol. 219, 17–24 (2018).

Article 
CAS 
PubMed 

Google Scholar
 

Kannan, N., Taylor, S. S., Zhai, Y., Venter, J. C. & Manning, G. Structural and functional diversity of the microbial kinome. PLoS Biol. 5, e17 (2007).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Kornev, A. P., Haste, N. M., Taylor, S. S. & Ten Eyck, L. F. Surface comparison of active and inactive protein kinases identifies a conserved activation mechanism. Proc. Natl Acad. Sci. USA 103, 17783–17788 (2006).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Klymiuk, V., Coaker, G., Fahima, T. & Pozniak, C. J. Tandem protein kinases emerge as new regulators of plant immunity. Mol. Plant Microbe Interact. 34, 1094–1102 (2021).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Yu, G. et al. The wheat stem rust resistance gene Sr43 encodes an unusual protein kinase. Nat. Genet. 55, 921–926 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Sung, Y. C. et al. Wheat tandem kinase RWT4 directly binds a fungal effector to activate defense. Nat. Genet. 57, 1238–1249 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Yu, G. et al. Aegilops sharonensis genome-assisted identification of stem rust resistance gene Sr62. Nat. Commun. 13, 1607 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Inoue, Y. et al. Evolution of the wheat blast fungus through functional losses in a host specificity determinant. Science 357, 80–83 (2017).

Article 
CAS 
PubMed 

Google Scholar
 

Sugihara, Y. et al. Disentangling the complex gene interaction networks between rice and the blast fungus identifies a new pathogen effector. PLoS Biol. 21, e3001945 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Guo, G. et al. The wheat NLR pair RXL/Pm5e confers resistance to powdery mildew. Plant Biotechnol. J. 23, 1260–1276 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Klymiuk, V. et al. Coordinated function of paired NLRs confers Yr84-mediated stripe rust resistance in wheat. Nat. Genet. 57, 1535–1542 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Gill, U. et al. Molecular and genetic characterization of barley mutants and genetic mapping of mutant rpr2 required for Rpg1-mediated resistance against stem rust. Theor. Appl. Genet. 129, 1519–1529 (2016).

Article 
CAS 
PubMed 

Google Scholar
 

Zhang, L. et al. Rpr1, a gene required for Rpg1-dependent resistance to stem rust in barley. Theor. Appl. Genet. 113, 847–855 (2006).

Article 
CAS 
PubMed 

Google Scholar
 

Running, K. L. D. et al. Development of diagnostic markers for the disease susceptibility gene Tsn1 in wheat reveals novel resistance alleles and a new locus required for ToxA sensitivity. Theor. Appl. Genet. 138, 164 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Dagvadorj, B., Outram, M. A., Williams, S. J. & Solomon, P. S. The necrotrophic effector ToxA from Parastagonospora nodorum interacts with wheat NHL proteins to facilitate Tsn1-mediated necrosis. Plant J. 110, 407–418 (2022).

Article 
CAS 
PubMed 

Google Scholar
 

Bernasconi, Z. et al. Virulence on Pm4 kinase-based resistance is determined by two divergent wheat powdery mildew effectors. Nat. Plants 12, 164–178 (2026).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Bernasconi, Z. et al. An HMA-like integrated domain in the wheat tandem kinase WTK4 recognises an RNase-like pathogen effector. Preprint at bioRxiv 10.1101/2025.08.26.672365 (2025).

Brueggeman, R. et al. The stem rust resistance gene Rpg5 encodes a protein with nucleotide-binding-site, leucine-rich, and protein kinase domains. Proc. Natl Acad. Sci. USA 105, 14970–14975 (2008).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Gaurav, K. et al. Population genomic analysis of Aegilops tauschii identifies targets for bread wheat improvement. Nat. Biotechnol. 40, 422–431 (2022).

Article 
CAS 
PubMed 

Google Scholar
 

Navathe, S. et al. ToxA–Tsn1 interaction for spot blotch susceptibility in Indian wheat: an example of inverse gene-for-gene relationship. Plant Dis. 104, 71–81 (2020).

Article 
CAS 
PubMed 

Google Scholar
 

Steuernagel, B. et al. The NLR-Annotator tool enables annotation of the intracellular immune receptor repertoire. Plant Physiol. 183, 468–482 (2020).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Cavalet-Giorsa, E. et al. Origin and evolution of the bread wheat D genome. Nature 633, 848–855 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Yan, X. et al. Molecular basis of SAP05-mediated ubiquitin-independent proteasomal degradation of transcription factors. Nat. Commun. 15, 1170 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Wang, Y. et al. An unusual tandem kinase fusion protein confers leaf rust resistance in wheat. Nat. Genet. 55, 914–920 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Li, M. et al. A membrane associated tandem kinase from wild emmer wheat confers broad-spectrum resistance to powdery mildew. Nat. Commun. 15, 3124 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Brueggeman, R. et al. The barley stem rust-resistance gene Rpg1 is a novel disease-resistance gene with homology to receptor kinases. Proc. Natl Acad. Sci. USA 99, 9328–9333 (2002).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Bender, K. W. et al. Activation loop phosphorylaton of a non-RD receptor kinase initiates plant innate immune signaling. Proc. Natl Acad. Sci. USA 118, e2108242118 (2021).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Guzmán-Vega, F. J. & Arold, S. T. AlphaCRV: a pipeline for identifying accurate binder topologies in mass-modeling with AlphaFold. Bioinform. Adv. 4, vbae131 (2024).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Homma, F., Huang, J. & van der Hoorn, R. A. L. AlphaFold-Multimer predicts cross-kingdom interactions at the plant–pathogen interface. Nat. Commun. 14, 6040 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Homma, F., Lyu, J. & van der Hoorn, R. A. L. Using AlphaFold Multimer to discover interkingdom protein–protein interactions. Plant J. 120, 19–28 (2024).

Article 
CAS 
PubMed 

Google Scholar
 

Dodds, P. N., Lawrence, G. J. & Ellis, J. G. Six amino acid changes confined to the leucine-rich repeat β-strand/β-turn motif determine the difference between the P and P2 rust resistance specificities in flax. Plant Cell 13, 163–178 (2001).

CAS 
PubMed 
PubMed Central 

Google Scholar
 

Farnham, G. & Baulcombe, D. C. Artificial evolution extends the spectrum of viruses that are targeted by a disease-resistance gene from potato. Proc. Natl Acad. Sci. USA 103, 18828–18833 (2006).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

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. Mol. Plant-Microbe Interact. 36, 434–446 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Seong, K. et al. Resurrection of the plant immune receptor Sr50 to overcome pathogen immune evasion. Preprint at bioRxiv https://doi.org/10.1101/2024.08.07.607039 (2025).