[1] Univ Bordeaux 1, Univ Bordeaux 2, UMR 1287, INRA, F-33883 Villenave Dornon, France
[2] Ecole Natl Ingn Travaux Agr, Unite Mixte Rech Sante Vegetale, INRA, F-33883 Villenave Dornon, France
[3] INRA, UMR 990, Inst Natl Polytechn, Ecole Natl Super Agron Toulouse, F-31326 Castanet Tolosan, France
[4] ALTADIS, Inst Tabac, F-24100 Bergerac, France
[5] Univ Bourgogne, UMR Plante Microbe Environm INRA 1088, CNRS 5184, F-21065 Dijon, France