全 文 :书西北植物学报!
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!"#*"-*"# !! 基金项目十二五(农村领域国家科技计划"
!"##22#""!"-"#!现代农业产业技术体系建设专项" 3245*%! # !! 作者简介贾彩红"
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条乙醇脱氢酶基因的克隆及表达分析
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