Rice is one of the world’s most important foods, but its production suffers from insect pests, causing losses of billions of dollars, and extensive use of environmentally damaging pesticides for their control1,2. However, the molecular mechanisms of insect resistance remain elusive. Although a few resistance genes for planthopper have been cloned, no rice germplasm is resistant to stem borers. Here, we report that biosynthesis of serotonin, a neurotransmitter in mammals3, is induced by insect infestation in rice, and its suppression confers resistance to planthoppers and stem borers, the two most destructive pests of rice2. Serotonin and salicylic acid derive from chorismate4. In rice, the cytochrome P450 gene CYP71A1 encodes tryptamine 5-hydroxylase, which catalyses conversion of tryptamine to serotonin5. In susceptible wild-type rice, planthopper feeding induces biosynthesis of serotonin and salicylic acid, whereas in mutants with an inactivated CYP71A1 gene, no serotonin is produced, salicylic acid levels are higher and plants are more insect resistant. The addition of serotonin to the resistant rice mutant and other brown planthopper-resistant genotypes results in a loss of insect resistance. Similarly, serotonin supplementation in artificial diet enhances the performance of both insects. These insights demonstrate that regulation of serotonin biosynthesis plays an important role in defence, and may prove valuable for breeding insect-resistant cultivars of rice and other cereal crops.

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  1. 1.

    Fahad, S. et al. Rice pest management and biological control. Sustain. Agric. Rev. 16, 85–106 (2015).

  2. 2.

    Chen, M., Shelton, A. & Ye, G. Y. Insect-resistant genetically modified rice in China: from research to commercialization. Annu. Rev. Entomol. 56, 81–101 (2011).

  3. 3.

    Berger, M., Gray, J. A. & Roth, B. L. The expanded biology of serotonin. Ann. Rev. Med. 60, 355–366 (2009).

  4. 4.

    Maeda, S. & Dudareva, N. The shikimate pathway and aromatic amino acid biosynthesis in plants. Annu. Rev. Plant Biol. 63, 73–105 (2012).

  5. 5.

    Fujiwara, T. et al. Sekiguchi lesion gene encodes a cytochrome P450 monooxygenase that catalyzes conversion of tryptamine to serotonin in rice. J. Biol. Chem. 285, 11308–11313 (2010).

  6. 6.

    Cheng, X., Zhu, L. & He, G. Towards understanding of molecular interactions between rice and the brown planthopper. Mol. Plant. 6, 621–634 (2013).

  7. 7.

    Du, B. et al. Identification and characterization of Bph14, a gene conferring resistance to brown planthopper in rice. Proc. Natl Acad. Sci. USA 106, 22163–22168 (2009).

  8. 8.

    Liu, Y. Q. et al. A gene cluster encoding lectin receptor kinases confers broad-spectrum and durable insect resistance in rice. Nat. Biotechnol. 33, 301–307 (2014).

  9. 9.

    Zhao, Y. et al. Allelic diversity in an NLR gene BPH9 enables rice to combat planthopper variation. Proc. Natl Acad. Sci. USA 113, 12850–12855 (2016).

  10. 10.

    Li, C. et al. Gene expression and plant hormone levels in two contrasting rice genotypes responding to brown planthopper infestation. BMC Plant Biol. 17, 57 (2017).

  11. 11.

    Qi, Y. X. et al. Serotonin modulates insect hemocyte phagocytosis via two different serotonin receptors. eLife 5, e12241 (2016).

  12. 12.

    Zhang, X. N. & Gaudry, Q. Functional integration of a serotonergic neuron in the Drosophila antennal lobe. eLife 5, e16836 (2016).

  13. 13.

    Erland, L. A. E., Turi, C. E. & Saxena, P. K. Serotonin: An ancient molecule and an important regulator of plant processes. Biotechnol. Adv. 34, 1347–1361 (2016).

  14. 14.

    Ueno, M., Imaoka, A., Kihara, J. & Arase, S. Tryptamine pathway-mediated DNA fragmentation is involved in sekiguchi lesion formation for light-emhanced resistance in lesion mimic mutant of rice to Magnaporthe grisea infection. J. Phytopathol. 156, 715–724 (2008).

  15. 15.

    Ishihara, A. et al. The tryptophan pathway is involved in the defense responses of rice against pathogenic infection via serotonin production. Plant J. 54, 481–495 (2008).

  16. 16.

    Ishihara, A., Hashimoto, Y., Miyagawa, H. & Wakasa, K. Induction of serotonin accumulation by feeding of rice striped stem borer in rice leaves. Plant Signal. Behav. 3, 714–716 (2008).

  17. 17.

    Xue, J. et al. Genome of rice pest brown planthopper and its endosymbionts reveal complex complementary contributions for host adaption. Genome Biol. 15, 521 (2014).

  18. 18.

    Arase, S. et al. Light-dependent accumulation of tryptamine in the rice sekiguchi lesion mutant infected with Magnaporthe grisea. J. Phytopathol. 149, 409–413 (2001).

  19. 19.

    Park, S., Lee, K., Kim, Y. S. & Back, K. Tryptamine 5-hydroxylase-deficient sekiguchi rice induces synthesis of 5-hydroxytryptophan and N- acetyltryptamine but decreases melatonin biosynthesis during senescence process of detached leaves. J. Pineal Res. 52, 211–216 (2012).

  20. 20.

    Tabashnik, B.E. & CarriŠre, Y. Surge in insect resistance to transgenic crops and prospects for sustainability. Nat. Biotechnol. 35, 926–935 (2017).

  21. 21.

    Lu, H. P. et al. Identification and characterization of a novel lesion mimic mutant in rice. J. Nucl. Agr. Sci. (Chin.) 30, 1037–1044 (2016).

  22. 22.

    Xu, R. F. et al. Gene targeting using the Agrobacterium tumefaciens-mediated CRISPR-Cas system in rice. Rice 7, 5 (2014).

  23. 23.

    Xu, G. et al. Identification and expression profiles of neuropeptides and their G protein-coupled receptors in the rice stem borer Chilo suppressalis. Sci. Rep. 6, 28976 (2016).

  24. 24.

    Pathak, P. K., Saxena, R. C. & Heinrichs, E. A. Parafilm sachet for measuring honeydew excretion by Nilaparvata lugens on Rice. J. Econ. Entomol. 75, 2 (1982).

  25. 25.

    Lou, Y. G. & Balswin, I. T. Manduca sexta recognition and resistance among allopolyploid Nicotiana host plants. Proc. Natl Acad. Sci. USA 100, 14581–14586 (2003).

  26. 26.

    Kang, S., Kang, K., Lee, K. & Back, K. Characterization of tryptamine 5-hydroxylase and serotonin synthesis in rice plants. Plant Cell Rep. 26, 2009–2015 (2007).

  27. 27.

    Ma, Z. Y., Guo, W., Guo, X. J., Wang, X. H. & Kang, L. Modulation of behavioral phase changes of the migratory locust by the catecholamine metabolic pathway. Proc. Natl Acad. Sci. USA 108, 3882–3887 (2011).

  28. 28.

    Cai, S. Y. et al. HsfA1a upregulates melatonin biosynthesis to confer cadium tolerance in tomato plants. J. Pineal Res. 62, e12387 (2017).

  29. 29.

    Ji, R. et al. A salivary endo-β-1,4-glucanase acts as an effector that enables the brown planthopper to feed on rice. Plant Physiol. 173, 1920–1932 (2017).

  30. 30.

    Han, L. Z., Li, S. B., Liu, P. L., Peng, Y. F. & Hou, M. L. New artificial diet for continuous rearing of Chilo suppressalis (Lepidoptera: Crambidae). Arthropod Biol. 105, 253–258 (2012).

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This study was supported by grants from National Key Research and Development Programme of China (2016YFD0102103), Agro-scientific Research in the Public Interest (201403030), Zhejiang Provincial S & T Project on Breeding of Agricultural (Food) Crops (2016C02050-2), China Postdoctoral Research Project (2017M620248), Dabeinong Funds for Discipline Development and Talent Training in Zhejiang University, China National Key Laboratory of Rice Biology, and the 111 Project. Assistance for melatonin measurement from J. Yu’s group is appreciated. We are grateful to T. Mou and Q. Fu for provision of BPH-resistant genotypes.

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Author notes

  1. These authors contributed equally: Hai-ping Lu, Ting Luo, Hao-wei Fu.


  1. State Key Laboratory of Rice Biology, Zhejiang Provincial Key Laboratory of Crop Genetic Resources, Institute of Crop Science, Zhejiang University, Hangzhou, China

    • Hai-ping Lu
    • , Yuan-yuan Tan
    • , Jian-zhong Huang
    •  & Qing-yao Shu
  2. State Key Laboratory of Rice Biology & Ministry of Agriculture Key Lab of Molecular Biology of Crop Pathogens and Insects, Institute of Insect Sciences, Zhejiang University, Hangzhou, China

    • Ting Luo
    • , Long Wang
    • , Gong-yin Ye
    •  & Yong-gen Lou
  3. Jiaxing Academy of Agricultural Sciences, Zhejiang, China

    • Hao-wei Fu
  4. Wuxi Hupper Bioseed Ltd., Wuxi, Jiangsu, China

    • Qing Wang
  5. School of Biology, Newcastle University, Newcastle upon Tyne, UK

    • Angharad M. R. Gatehouse
  6. Hubei Collaborative Innovation Center for Grain Industry, Jingzhou, Hubei, China

    • Qing-yao Shu


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Q.Y.S., Y.G.L., A.M.R.G., G.Y.Y. and J.Z.H. contributed to study design and data analysis, H.L. contributed to overall study and data analysis, T.L. contributed to BPH and WBPH resistance studies, H.W.F. contributed to Jiazhe LM mutant development and field studies, L.W. contributed SSB resistance studies, Q.W. and Y.Y.T. contributed to the development and characterization of knockout mutants, and A.M.R.G. and Q.Y.S. wrote the manuscript. All authors read and approve the paper.

Competing interests

The authors declare no competing interests.

Corresponding authors

Correspondence to Angharad M. R. Gatehouse or Yong-gen Lou or Qing-yao Shu.

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