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Sichuan Univ, Coll Life Sci, Key Lab Bioresource & Ecoenvironm, Minist Educ,Sichuan Key Lab Mol Biol & Biotechnol, Chengdu 610065, Peoples R China Sichuan Univ, Coll Life Sci, Key Lab Bioresource & Ecoenvironm, Minist Educ,Sichuan Key Lab Mol Biol & Biotechnol, Chengdu 610065, Peoples R China