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S China Agr Univ, Coll Life Sci, Key Lab, Minist Educ Rice Fertil Dev & Stress Resistance, Guangzhou 510642, Guangdong, Peoples R China S China Agr Univ, Coll Life Sci, Key Lab, Minist Educ Rice Fertil Dev & Stress Resistance, Guangzhou 510642, Guangdong, Peoples R China