Sub1A is an ethylene-response-factor-like gene that confers submergence tolerance to rice

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Most Oryza sativa cultivars die within a week of complete submergence—a major constraint to rice production in south and southeast Asia that causes annual losses of over US$1 billion and affects disproportionately the poorest farmers in the world1,2. A few cultivars, such as the O. sativa ssp. indica cultivar FR13A, are highly tolerant and survive up to two weeks of complete submergence owing to a major quantitative trait locus designated Submergence 1 (Sub1) near the centromere of chromosome 9 (refs 3, 4, 5–6). Here we describe the identification of a cluster of three genes at the Sub1 locus, encoding putative ethylene response factors. Two of these genes, Sub1B and Sub1C, are invariably present in the Sub1 region of all rice accessions analysed. In contrast, the presence of Sub1A is variable. A survey identified two alleles within those indica varieties that possess this gene: a tolerance-specific allele named Sub1A-1 and an intolerance-specific allele named Sub1A-2. Overexpression of Sub1A-1 in a submergence-intolerant O. sativa ssp. japonica conferred enhanced tolerance to the plants, downregulation of Sub1C and upregulation of Alcohol dehydrogenase 1 (Adh1), indicating that Sub1A-1 is a primary determinant of submergence tolerance. The FR13A Sub1 locus was introgressed into a widely grown Asian rice cultivar using marker-assisted selection. The new variety maintains the high yield and other agronomic properties of the recurrent parent and is tolerant to submergence. Cultivation of this variety is expected to provide protection against damaging floods and increase crop security for farmers.

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Figure 1: Sub1 region gene composition and submergence-induced mRNA accumulation in rice.
Figure 2: Characterization of submergence response in transgenic rice ectopically expressing Sub1A-1.
Figure 3: Introgression of the FR13A Sub1 haplotype into an intolerant variety by MAS confers submergence tolerance.


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We thank J. Dvorak for discussions of experimental design, H. Zhang for providing BAC clones, D. Chen, J. S. Jeon, J. Ni, P. Colowit, D. Ruan, R. Bruskiewich, G. Vergara, M. Nas, A. Pamplona, C. N. Neeraja, and S. Singh for their technical assistance and suggestions, support and discussions, and the IRRI Grain Quality, Nutrition, and Postharvest Center for amylose data. This work was supported by USDA-NRICGP grants to D.J.M. and P.C.R., and to J.B.-S. and P.C.R., and USAID Linkage Project funds to J.B.-S. and P.C.R. The work on transferring Sub1 to Swarna was supported by a grant from the German Federal Ministry for Economic Cooperation and Development (BMZ) to D.J.M. and A.M.I.

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Correspondence to Pamela C. Ronald or David J. Mackill.

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Sequences were submitted to GenBank/EMBL/DDJB under accession numbers DQ011597–DQ011607 and DQ453964–DQ453966. Reprints and permissions information is available at The authors declare no competing financial interests.

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