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INRA, Ctr Versailles Grignon, AgroParisTech, Microbiol & Mol Genet Lab,CNRS,UMR2585,UMR1238, F-78850 Thiverval Grignon, France INRA, Ctr Versailles Grignon, AgroParisTech, Microbiol & Mol Genet Lab,CNRS,UMR2585,UMR1238, F-78850 Thiverval Grignon, France