[1] Univ Clermont Auvergne, INRA, UMR GDEC, Clermont Ferrand, France
[2] Clermont Auvergne Univ, Genet Divers & Ecophysiol Cereals, UMR1095, Clermont Ferrand, France
[3] Univ Paris Sud, Univ Paris Saclay, CNRS, UMR GQE,INRA,Agro ParisTech, Gif Sur Yvette, France
[4] Univ Toulouse, CNRS, UMR5219, Inst Math Toulouse, Toulouse, France
[5] Univ Calif Riverside, Dept Bot & Plant Sci, Riverside, CA 92521 USA
[6] Univ Montpellier, Montpellier SupAgro, INRA, LEPSE, Montpellier, France