Huang Y, Yin C, Liu J, Feng B, Ge D, Kong L, Ortiz-Morea FA, Richter J, Hauser MT, Wang WM, Shan L, He P. A trimeric CrRLK1L-LLG1 complex genetically modulates SUMM2-mediated autoimmunity. Nature Communications. 2020 Sep 25;11(1):4859.

Liu J, Huang Y, Kong L, Yu X, Feng B, Liu D, Zhao B, Mendes GC, Yuan P, Ge D, Wang WM, Fontes EPB, Li P, Shan L, He P. The malectin-like receptor-like kinase LETUM1 modulates NLR protein SUMM2 activation via MEKK2 scaffolding. Nature Plants. 2020 Sep;6(9):1106-1115.

Ma X, Claus LAN, Leslie ME, Tao K, Wu Z, Liu J, Yu X, Li B, Zhou J, Savatin DV, Peng J, Tyler BM, Heese A, Russinova E, He P, Shan L. Ligand-induced monoubiquitination of BIK1 regulates plant immunity. Nature. 2020 May;581(7807):199-203.

Liu J, Li J, Shan L. SERKs. Current Biology. 2020 Apr 6;30(7):R293-4.

Yang Y, Liu J, Yin C, de Souza Vespoli L, Ge D, Huang Y, Feng B, Xu G, Manhães AMEA, Dou S, Criswell C, Shan L, Wang X, He P. RNA Interference-Based Screen Reveals Concerted Functions of MEKK2 and CRCK3 in Plant Cell Death Regulation. Plant Physiology. 2020 May;183(1):331-344.

Levy JG, Gross R, Mendoza-Herrera A, Tang X, Babilonia K, Shan L, Kuhl JC, Dibble MS, Xiao F, Tamborindeguy C. Lso-HPE1, an Effector of ‘Candidatus Liberibacter solanacearum’, Can Repress Plant Immune Response. Phytopathology. 2020 Mar;110(3):648-655.



Cox, K. L., Babilonia, K., He, P., and Shan, L. (2019) Return of old foes – recurrence of bacterial blight and fusarium wilt of cotton. Current Opinion in Plant Biology 50: 95-103.

Xiong, C., Luo, D., Lin, A., Zhang, C., Shan, L., He, P., Li, B., Zhang, Q., Hua, B., Yuan, Z., Li, H., Zhang, J., Yang, C., Lu, Y., Ye, Z., and Wang, T. (2019) A tomato B-box protein SlBBX20 modulates carotenoid biosynthesis by directly activating PHYTOENE SYNTHASE 1, and is targeted for 26S proteasome-mediated degradation. New Phytologist 221: 279–294.

Chen, C., Yang, S., Liu, Y., Jamieson, P., Shan, L., and Chu, K. (2019) Accumulation and phytotoxicity of perfluorooctanoic acid and 2,3,3,3-tetrafluoro-2-(heptafluoropropoxy)propanoate in Arabidopsis thaliana and Nicotiana benthamiana. Environmental Pollution 259:113817

Li B, Ferreira MA, Huang M, Camargos LF, Yu X, Teixeira RM, Carpinetti PA, Mendes GC, Gouveia-Mageste BC, Liu C, Pontes CSL, Brustolini OJB, Martins LGC, Melo BP, Duarte CEM, Shan L, He P, Fontes EPB. The receptor-like kinase NIK1 targets FLS2/BAK1 immune complex and inversely modulates antiviral and antibacterial immunity. Nature Communications. 2019 Nov 1;10(1):4996.

Yu X, Xu G, Li B, de Souza Vespoli L, Liu H, Moeder W, Chen S, de Oliveira MVV, Ariádina de Souza S, Shao W, Rodrigues B, Ma Y, Chhajed S, Xue S, Berkowitz GA, Yoshioka K, He P, Shan L. The Receptor Kinases BAK1/SERK4 Regulate Ca2+ Channel-Mediated Cellular Homeostasis for Cell Death Containment. Current Biology. 2019 Nov 18;29(22):3778-3790.e8.

Yu X, Li B, Jang GJ, Jiang S, Jiang D, Jang JC, Wu SH, Shan L, He P. Orchestration of processing body dynamics and mRNA decay in Arabidopsis immunity. Cell reports. 2019 Aug 20;28(8):2194-205.

Mu C, Zhou L, Shan L, Li F, Li Z. Phosphatase GhDs PTP 3a interacts with annexin protein Gh ANN 8b to reversely regulate salt tolerance in cotton (Gossypium spp.). New Phytologist. 2019 Sep;223(4):1856-72.

Zhou J, Wang P, Claus LA, Savatin DV, Xu G, Wu S, Meng X, Russinova E, He P, Shan L. Proteolytic processing of SERK3/BAK1 regulates plant immunity, development, and cell death. Plant physiology. 2019 May 1;180(1):543-58.

Levy, Julien Gad, Rachel Gross, M. Azucena Mendoza Herrera, Xiaotian Tang, Kevin Babilonia, Libo Shan, Joseph Kuhl, Maggie Dibble, Fangming Xiao, and Cecilia Tamborindeguy. “Lso-HPE1, an effector of ‘Candidatus Liberibacter solanacearum’can repress plant immune response.” Phytopathology ja (2019).

Wan, W., Zhang, L., Pruitt, R., Zaidem, M., Brugman, R., Ma, X., Krol, E., Perraki, A., Kilian, J., Grossmann, G., Stahl, M., Shan, L., Zipfel, C., van Kan, J., Hedrich, R., Weigel, D., Gust, A., and Nuernberger, T. Comparing Arabidopsis receptor kinase and receptor protein-mediated immune signaling reveals BIK1-dependent differences. New Phytologist (2018).

Shan, L., and He, P. (2018) Pipped at the post: pipecolic acid derivative identified as SAR regulator. Cell 173:286-287.

Jamieson, P.A., Shan, L., and He, P. Plant cell surface molecular cypher: receptor-like proteins and their roles in immunity and development. Plant Science 274:242-251.

Lal, N.K., Nagalakshmi, U., Hurlburt, N.K., Flores, R., Bak, A., Sone, P., Ma, X., Song, G., Walley, J., Shan, L., He, P., Casteel, C., Andrew J. Fisher, A.J., and Dinesh-Kumar, S.P. (2018) Nuclear role of a plant receptor-like cytoplasmic kinase in hormone regulation during innate immunity. Cell Host & Microbe 23:485-497.

Zhou, J., Liu, D., Wang, P., Lin, W., Chen, S., Mishev, K., Lu, D., Kumar, R., Vanhoutte, I., He, P., Russinova, E., and Shan, L. (2018) Regulation of Arabidopsis brassinosteroid receptor BRI1 endocytosis and degradation by plant U-box PUB12/PUB13-mediated ubiquitination. Proc Natl Acad Sci U S A. 115: E1906-E1915

Huang, Y., Jamieson, P., and Shan, L. (2018) The APEX approaches: a unified LRR-RK network revealed. Trends in Plant Science 23: 372-374. .

Fan, J., Bai, P., Xiong, Y., Ning, Y., Zhang, C., Wang, R., Meng, X., Zhou, J., Wang, M., Park, C.H., Bellizzi, M., Jeon, J.S. Shan, L., and Guo-Liang Wang (2018) The monocot-specific receptor-like kinase SDS2 modulated by the E3 Ligase SPL11 controls cell death and immunity via receptor-like cytoplasmic kinases in rice. Cell Host & Microbe.

He Y., Zhou J., Shan L., and Meng X. (2018) Plant cell surface receptor-mediated signaling – a common theme amid diversity. J Cell Sci. 131(2).

Li, F., Li, M., Wang, P. Cox, K. L., Duan, L., Dever, J. K., Shan, L., Li, Z., He, P. (2017). Regulation of cotton (Gossypium hirsutum) drought responses by mitogen‐activated protein (MAP) kinase cascade‐mediated phosphorylation of GhWRKY59. New Phytologist 215: 1462-1475.

Cox, K. L., Meng, F., Wilkins, K. E., Li, F., Wang, P., Booher, N. J., Carpenter, S. C. D., Chen, L. Q., Zheng, H., Gao, X., Zheng, Y., Fei, Z., Yu, J. Z., Isakeit, T., Wheeler, T., Frommer, W. B., He, P., Bogdanove, A. J., and Shan, L. (2017) TAL effector driven induction of a SWEET gene confers susceptibility to bacterial blight of cotton. Nature Communications.

Mang H., Feng, B., Hu, Z., Boisson-Dernier, A., Franck, C.M., Meng, X., Xu, G., Wang, T., Shan, L., and He, P. (2017) Differential regulation of two-tiered plant immunity and sexual reproduction by ANXUR receptor-like kinases. The Plant Cell 29: 3140–3156.

Feng, B., Ma, S., Chen, S., Zhu, N., Zhang, S., Yu, B., Yu, Y., Le, B., Chen, X., Dinesh-Kumar, S.P., Shan, L., and He, P. (2016) PARylation of the forkhead-associated domain protein DAWDLE regulates plant immunity. EMBO Reports 17:1799-1813.

Feng, B., Liu, C., Shan, L., and He, P. (2016) ADP-ribosylation takes control in plant-bacterium interactions. PLoS Pathog.

Ma, X., Xu, G., He, P., and Shan, L. (2016) SERKing Coreceptors for Receptors. Trends Plant Sci.

Liu, Y., Huang, X., Li, M., He, P., and Zhang, Y. (2016) Loss-of-function of Arabidopsis receptor-like kinase BIR1 activates cell death and defense responses mediated by BAK1 and SOBIR1. New Phytologist.

Yadeta, K. A., Elmore, J. A., Creer, A.Y., Feng, B., Franco, J. Y., Rufian, J. S., He, P., Phinney, B., and Coaker, G. (2016) Functional analyses of extracellular cysteine residues and phosphorylation for immune responses mediated by the Arabidopsis cysteine-rich receptor-like kinase CRK28. Plant Physiology.

Eschen-Lippold, L., Jiang, X., Elmore, J.M., Mackey, D., Shan, L., Coaker, G., Scheel, D., and Lee, J. (2016). Bacterial AvrRpt2-Like Cysteine Proteases Block Activation of the Arabidopsis Mitogen-Activated Protein Kinases, MPK4 and MPK11. Plant Physiology.

Li, B., Meng, X., Shan L., and He, P. (2016) Transcriptional Regulation of Pattern-Triggered Immunity in Plants. Cell Host & Microbe.

Oliveira, M.V.V., Xu, G., Meng, X., Chen, X., Intorne, A.C., Babinat, A., Filho, G.A.S., Shan, L., and He, P. (2016) Specific control of Arabidopsis BAK1- and SERK4-regulated cell death by protein glycosylation. Nature Plants.

Meng, X., Zhou, J., Tang, J., Li, B., Oliveira, M.V.V., Chai., J., He, P. and Shan, L. (2016) Ligand-induced receptor-like kinase complex regulates floral organ abscission in Arabidopsis. Cell Reports 14, 1330–1338.

Meng, X., Shan, L., and He, P. (2015) Stack heterotrimeric G proteins and MAPK cascades on a RACK. Molecular Plant 8:1691-1693.

Meng, X., Chen, X., Mang, H., Liu, C., Yu, X., Gao, X., Torii, K., He, P., and Shan,. L. (2015) Differential Function of Arabidopsis SERK Family Receptor-like Kinases in Stomatal Patterning. Current Biology. DOI:

Zhou, J., Lu, D., Xu, G., Finlayson. SA., He, P., and Shan, L. (2015) The dominant negative ARM domain uncovers multiple functions of PUB13 in Arabidopsis immunity, flowering, and senescence. J Exp Bot.

Li, B., Jiang, S., Yu, X., Cheng, C., Chen, S., Cheng, Y., Yuan, JS, Jiang, D., He, P., and Shan, L. (2015) Phosphorylation of Trihelix Transcriptional Repressor ASR3 by MAP KINASE4 Negatively Regulates Arabidopsis Immunity. Plant Cell.

Feng, B., Liu, C., de Oliveira MV, Intorne AC, Li, B., Babilonia, K., de Souza Filho G.A., Shan, L., and He, P. (2015) Protein Poly(ADP-ribosyl)ation Regulates Arabidopsis Immune Gene Expression and Defense Responses. PLoS Genetics.



Li, F., Cheng, C., Cui, F., Oliveira, M.V.V., Yu, X., Meng, X., Intorne, A.C., Babilonia, K., Li, M., Chen, X., Ma, X., Xiao, S., Zeng, Y., Fei, Z., Metz, R., Johnson, C.D., Koiwa, H., Sun, W., Li, Z., Filho, G.A.S., Shan, L., and He, P. (2014) Modulation of RNA polymerase II phosphorylation downstream of pathogen perception orchestrates plant immunity. Cell Host & Microbe 16(6):748-58.

Feng, B., and Shan, L., (2014) ROS open roads to roundworm infection. Science Signaling 7(320), pe10. 

Wu, S., Shan, L., and He, P. (2014) Microbial signature-triggered plant defense responses and early signaling mechanisms. Plant Science 228:118-126.

Gao, X., Cox Jr., K.L., and He, P. (2014) Functions of calcium-dependent protein kinases in plant innate immunity. Plants 3:160-176.

Lin, W., Li, B., Lu, D., Chen, S., Zhu, N., He, P., and Shan L. (2014) Tyrosine phosphorylation of BAK1/BIK1 mediates Arabidopsis innate immunity. Proc Natl Acad Sci U S A. 111: 3632-3637

Zhou, J., Wu, S., Chen, X., Liu, C., Sheen, J., Shan, L., and He, P. (2014). Pseudomonas syringae effector HopF2 suppresses Arabidopsis immunity by targeting BAK1. The Plant Journal 77:235–245.

Zhou, J, He, P., and Shan, L. Ubiquitination of plant immune receptors. (2014) Methods in Mol Biol. 1209:219-231. doi: 10.1007/978-1-4939-1420-3_17.

Cheng, C, Gao, X, Feng, B, Sheen, J, Shan, L, and He, P. (2013) Plant immune response to pathogens differs with changing temperatures. Nat Commun. 4:2530. doi: 10.1038/ncomms3530

Lin, W., Lu, D., Gao, X., Jian, S., Ma, X., Wang, Z., Mengiste, T., He, P., and Shan, L. (2013) Inverse modulation of plant immunity and plant development by a receptor-like cytoplasmic kinase BIK1. Proc Natl Acad Sci U S A. 110 (29):12114-9.

Cui, F., Wu, S., Sun, W., Coaker, G., Kunkel, B. N, He, P., and Shan, L. (2013) The Pseudomonas syringae type III effector AvrRpt2 promotes pathogen virulence via stimulating Arabidopsis Auxin/Indole Acetic Acid protein turnover. Plant Physiology 162:1018-1029.

Gao, X., Chen, X., Lin, W., Lu, D., Niu, Y., Li, L., Cheng, C., McCormack, M., Sheen, J., Shan, L., and He, P. (2013) Bifurcation of Arabidopsis NLR Immune Signaling via Ca2+-Sensor Protein Kinases. PLoS Pathog 9, e1003127.

Gao, X., and He, P. (2013) Nuclear dynamics of Arabidopsis calcium-dependent protein kinases in effector-triggered immunity. Plant Signaling & Behavior 8(4). pii: e23868.

Lin, W., Ma, X., Shan, L., and He, P. (2013) Big roles of small kinases: the complex functions of receptor-like cytoplasmic kinases in plant immunity and development. Journal of Integrative Plant Biology. 55:1188-1197

Gao, X., Li, F., Li, M., Kianinejad, A., Dever, J., Wheeler, T., Li, Z., He, P., and Shan, L. (2013) Cotton GhBAK1 mediates Verticillium wilt resistance and cell death. Journal of Integrative Plant Biology 55:586-596.

Lu, D., Lin, W., Gao, X., Wu, S., Cheng, C., Avila, J., Heese, A., Devarenne, T., He, P., and Shan, L. (2011) Direct ubiquitination of pattern recognition receptor FLS2 attenuates plant innate immunity. Science. 332: 1439-1442.

Gao, X., Britt, R.C., Shan, L. and He, P. (2011) Agrobacterium-mediated virus-induced gene silencing assay in cotton. J Vis Exp. 54.

Gao, X., Wheeler, T., Li, Z., Kenerley, C., He, P., and Shan, L. (2011) Silencing GhNDR1 and GhMKK2 compromised cotton resistance to Verticillium wilt. The Plant Journal. 66:293-305.

Wu, S., Lu, D., Kabbage, M., Wei, H., Swingle, B., Dickman, M., He, P., and Shan, L. (2011) Bacterial effector HopF2 suppresses Arabidopsis innate immunity at the plasma membrane. Mol. Plant Microbe Interact. 24:585-93.

Lu, D., Wu, S., Gao, X., Zhang, Y., Shan, L. and He, P. (2010) A receptor-like cytoplasmic kinase BIK1 associates with flagellin receptor complex to initiate plant innate immunity. Proc Natl Acad Sci U S A. 107:496-501.

Lu, D., He, P., and Shan, L. (2010) Bacterial effectors target BAK1-associated receptor complexes: one stone two birds. Communicative & Integrative Biology. 3 (2): 1-4.

Boudsocq, M., Willmann R., McCormack, M., Lee, H., Shan L., He P., Bush, J., Cheng, S., and Sheen J., (2010) Differential innate immune signalling via Ca2+ sensor protein kinases. Nature. 464: 418-422.


  • Chinchilla, D., Shan, L., He, P., de Vries, S. and Kemmerling, B. (2009) One for all-the receptor-associated kinase BAK1. Trend in Plant Science.14 (10) 535-541.
  • Shan, L., He, P., Li, J., Heese, A., Peck, S. C., Nurnberger, T., Martin, G. B., and Sheen, J. (2008). Bacterial effectors target the common signaling partner BAK1 to disrupt multiple MAMP receptor-signaling complexes and impede plant immunity. Cell Host Microbe 4, 17-27.
  • Shang, Y., Li, X., Cui, H., He, P., Thilmony, R., Chintamanani, S., Zwiesler-Vollick, J., Gopalan, S., Tang, X., and Zhou, J.-M. (2006) RAR1, a central player in plant immunity, is targeted by Pseudomonas syringae effector AvrB. Proc Natl Acad Sci U S A  103:19200-19205.
  • Xiao, F., He, P., Abramovitch, R., Dawson, J., Nicholson, L., Sheen, J., and Martin, G. (2007) The N-terminal region of Pseudomonas type III effector AvrPtoB elicits Pto-dependent immunity and has two distinct virulence determinants. The Plant Journal 52: 595-614.
  • Shan, L., He, P., and Sheen, J. (2007) Intercepting host MAPK signaling cascades by bacterial type III effectors. Cell Host Microbe 1: 167-174.
  • He, P., Shan, L., and Sheen, J. (2007) Elicitation and suppression of MAMP-triggered immunity in plant-microbe interactions. Cell Microbiol 9: 1385-1396.
  • Sheen, J., and He, P. (2007) Nuclear actions in innate immune signaling. Cell 128:821-823.
  • Shang, Y., Li, X., Cui, H., He, P., Thilmony, R., Chintamanani, S., Zwiesler-Vollick, J., Gopalan, S., Tang, X., and Zhou, J.-M. (2006) RAR1, a central player in plant immunity, is targeted by Pseudomonas syringae effector AvrB. Proc Natl Acad Sci U S A. 103:19200-19205.
  • He, P., Shan, L. and Sheen, J. (2006) The use of protoplasts to study innate immune responses. Methods Mol Biol. 354: 1-10.
  • He, P., Shan, L., Lin, N.-C., Martin, G., Kemmerling, B., Nurnberger, T., and Sheen, J. (2006). Specific bacterial suppressors of MAMP signaling upstream of MAPKKK in Arabidopsis innate immunity. Cell 125: 563-575.
  • Robert-Seilaniantz, A.*, Shan, L.*, Zhou, J.-M., and Tang, X. (2006). The Pseudomonas syringae tomato DC3000 type III effector HopF2 has a putative myristoylation site required for its avirulence and virulence functions. Mol. Plant Microbe Interact. 19: 130-138. (*Co-first author)


  • Shan, L., Oh, H., Chen, J., Guo, M., Zhou, J.-M., Alfano, J.R., Collmer, A., Jia, X., and Tang X. The HopPtoF Locus of Pseudomonas syringae pv. tomato DC3000 Encodes a Type III Chaperone and a Cognate Effector. Mol. Plant Microbe Interact. 17: 447-455.
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  • He, P., Friebe, B. R., Gill, B. S., and Zhou, J.-M. (2003). Allopolyploidy alters gene expression in the highly stable hexaploid wheat. Plant Molecular Biology 52: 401-414.
  • Li, J., Shan, L., Zhao, T., Zhou, J.-M., and Tang, X. (2002). Overexpression of Pto Induces a Salicylate-Independent Cell Death But Inhibits Necrotic Lesions Caused by Salicylate-Deficiency in Tomato Plants. Mol. Plant Microbe Interact. 15: 654-661.
  • Petnicki-Ocwieja, T., Schneider, D. J., Tam, V. C., Chancey, S. T., Shan, L., Jamir, Y., Schechter, L. M., Janes, M. D., Buell, C. R., Tang, X., et al. (2002).Genome wide identification of proteins secreted by the Hrp type III protein secretion system of Pseudomonas syringae pv. tomato DC3000. Proc. Natl. Acad. Sci. U S A 99: 7652-7657.
  • He, P., Warren, R. F., Zhao, T., Shan, L., Zhu, L., Tang, X., and Zhou, J.-M. (2001). Overexpression of Pti5 in tomato potentiates pathogen-induced defense gene expression and enhances disease resistance to Pseudomonas syringae pv. tomato. Molecular Plant-Microbe Interactions 14: 1453-1457.
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  • Gong, J., Zheng, X., Du, B., Qian, Q., Chen, S., Zhu, L. and He, P. (2001).Comparative study of QTLs for agronomic traits of rice (Oryza sativa) between salt stress and nonstress environment. Science in China (Series C) 44: 73-82.
  • Qian, Q., He, P., Teng, S., Zeng, D., and Zhu, L. (2001).QTLs analysis of tiller angle in rice (Oryza sativa L.) Acta Genetica Sinica 28: 29-32.
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  • Shan, L., Li, Y.W., Zhao, T.H., Liang, H., Ouyang, J.W., Jia, S.E., and Jia, X. (2000). Estimation of biolistic transformation effect by transient expression of C1-R regulatory genes of anthocyanin biosynthesis. Acta Genetica Sinica 27: 65-69.
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