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Many developmental processes in plants are regulated by GA hormones. GA homeostasis is achieved via complex biosynthetic and catabolic pathways. GA catabolic enzymes include GA 2-oxidases that are classified into three classes. Members of class III GA 2-oxidases typically act on GA precursors containing a C 20 -skeleton. Here, we identified two further members of this class of GA 2-oxidases, namely AtGA2ox9 and AtGA2ox10 , in the Arabidopsis ( Arabidopsis thaliana ) genome. Both genes encode enzymes that have functional similarities to AtGA2ox7 and AtGA2ox8, which are class III GA 2-oxidases that 2β-hydroxylate C 20 -GAs. Previously unknown for GA 2-oxidases, AtGA2ox9 performs 2α-hydroxylation of C 19 -GAs and harbors putative desaturating activity of C 20 -GAs. Additionally, AtGA2ox9 and AtGA2ox10 exhibit GA 20-oxidase activity. AtGA2ox9 oxidizes carbon-20 to form tricarboxylic acid C 20 -GAs, whereas AtGA2ox10 produces C 19 -GA 9  AtGA2ox9 transcript levels increase after cold treatment and AtGA2ox10 is expressed mainly in the siliques of Arabidopsis plants. Atga2ox9 loss-of-function mutants are more sensitive to freezing temperatures, whereas Atga2ox10 loss-of-function mutants produce considerably more seeds per silique than wild-type plants. We conclude that in Arabidopsis, AtGA2ox9 contributes to freezing tolerance and AtGA2ox10 regulates seed production.

The Class III Gibberellin 2-Oxidases AtGA2ox9 and AtGA2ox10 Contribute to Cold Stress Tolerance and Fertility