Bph6 encodes an exocyst-localized protein and confers broad resistance to planthoppers in rice

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The brown planthopper (BPH) and white-backed planthopper (WBPH) are the most destructive insect pests of rice, and they pose serious threats to rice production throughout Asia. Thus, there are urgent needs to identify resistance-conferring genes and to breed planthopper-resistant rice varieties. Here we report the map-based cloning and functional analysis of Bph6, a gene that confers resistance to planthoppers in rice. Bph6 encodes a previously uncharacterized protein that localizes to exocysts and interacts with the exocyst subunit OsEXO70E1. Bph6 expression increases exocytosis and participates in cell wall maintenance and reinforcement. A coordinated cytokinin, salicylic acid and jasmonic acid signaling pathway is activated in Bph6-carrying plants, which display broad resistance to all tested BPH biotypes and to WBPH without sacrificing yield, as these plants were found to maintain a high level of performance in a field that was heavily infested with BPH. Our results suggest that a superior resistance gene that evolved long ago in a region where planthoppers are found year round could be very valuable for controlling agricultural insect pests.

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Fig. 1: Map-based cloning of Bph6.
Fig. 2: Molecular characterization of Bph6.
Fig. 3: Demonstration that BPH6 interacts with exocyst subunit OsEXO70E1.
Fig. 4: Characterization of exocyst gene expression and cell walls in 9311-Bph6-NIL and 9311 plants.
Fig. 5: Analyses of phytohormones in Bph6-carrying plants.
Fig. 6: Characterization of Bph6-mediated resistance.
Fig. 7: Evolution of Bph6 alleles in rice.


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We thank Q. Qian (China National Rice Research Institute), L. Han (Chinese Academy of Agricultural Sciences), L. Yan (Jiangxi Academy of Agricultural Sciences) and D. Pan (Guangdong Academy of Agricultural Sciences) for kindly providing rice germplasm, S. Wang (Huazhong Agricultural University) for kindly providing the rice disease pathogen PXO145, Y. Lin (Huazhong Agricultural University) for kindly providing the striped stem borer insects, D. Zeng (China National Rice Research Institute) for kindly providing the WBPH insects, Y. Liu for suggestions for the experiments, and Q. Zhang, R. He and J. Blackwell for edits and suggestions. This work was supported by grants from the National Natural Science Foundation of China (31230060 and 31630063, both to G.H.), the National Program on Research and Development of Transgenic Plants (2016ZX08009-003-001 to G.H.) and the National Key Research and Development Program (2016YFD0100600 and 2016YFD0100900, both to G.H.).

Author information

G.H. conceived and supervised the project; G.H. and J.G. designed the experiments; J.G. performed most of the experiments; C.X., Y.Z., D.W., B.D., X.W., Y.O., X.L., W.W., Y.Q., S.J., B.C., X.S., H.W., Y.M., Y.W., L.H., S.S., L.Z., X.X., R.C. and Y.F. performed some of the experiments; and J.G., C.X., D.W., B.D., Y.Z. and G.H. analyzed data and wrote the manuscript.

Correspondence to Bo Du or Guangcun He.

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Supplementary information

Supplementary Text and Figures

Supplementary Figures 1–27 and Supplementary Tables 1, 2, 4, 5, 7, 10 and 11

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Supplementary Table 3

GeneChip data for comparing Nip-Bph6-NIL and Bph6-RNAi plants

Supplementary Table 6

Information on accessions of wild rice and cultivated varieties that were sequenced in the Bph6 coding region

Supplementary Table 8

List of SNPs between Swarnalata or Nipponbare and the 80 haplotypes

Supplementary Table 9

List of indels between Swarnalata or Nipponbare and the 80 haplotypes

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