title: Photoperiod-H1 (Ppd-H1) Controls Leaf Size.
abstract: Leaf size is a major determinant of plant photosynthetic activity and biomass; however, it is poorly understood how leaf size is genetically controlled in cereal crop plants like barley (Hordeum vulgare). We conducted a genome-wide association scan for flowering time, leaf width, and leaf length in a diverse panel of European winter cultivars grown in the field and genotyped with a single-nucleotide polymorphism array. The genome-wide association scan identified PHOTOPERIOD-H1 (Ppd-H1) as a candidate gene underlying the major quantitative trait loci for flowering time and leaf size in the barley population. Microscopic phenotyping of three independent introgression lines confirmed the effect of Ppd-H1 on leaf size. Differences in the duration of leaf growth and consequent variation in leaf cell number were responsible for the leaf size differences between the Ppd-H1 variants. The Ppd-H1-dependent induction of the BARLEY MADS BOX genes BM3 and BM8 in the leaf correlated with reductions in leaf size and leaf number. Our results indicate that leaf size is controlled by the Ppd-H1- and photoperiod-dependent progression of plant development. The coordination of leaf growth with flowering may be part of a reproductive strategy to optimize resource allocation to the developing inflorescences and seeds.
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Benedikt
authors.initials: B
authors.lastname: Digel
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Elahe
authors.initials: E
authors.lastname: Tavakol
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Gabriele
authors.initials: G
authors.lastname: Verderio
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Alessandro
authors.identifier: 0000-0001-9299-7773
authors.initials: A
authors.lastname: Tondelli
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Xin
authors.initials: X
authors.lastname: Xu
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.).
authors.forename: Luigi
authors.identifier: 0000-0002-6067-4600
authors.initials: L
authors.lastname: Cattivelli
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.) laura.rossini@unimi.it korff@mpipz.mpg.de.
authors.forename: Laura
authors.identifier: 0000-0001-6509-9177
authors.initials: L
authors.lastname: Rossini
authors.affiliation: Max Planck Institute for Plant Breeding Research, D-50829 Cologne, Germany (B.D., M.v.K.);Institute of Plant Genetics, Heinrich-Heine-University, 40225 Duesseldorf, Germany (B.D., M.v.K.);Cluster of Excellence on Plant Sciences "From Complex Traits Towards Synthetic Modules," 40225 Duesseldorf, Germany (B.D., M.v.K.);Università degli Studi di Milano-DiSAA, 20133 Milan, Italy (E.T., G.V., L.R.);Department of Crop Production and Plant Breeding, College of Agriculture, Shiraz University, 7144165186 Shiraz, Iran (E.T.);Council for Agricultural Research and Economics, Genomics Research Centre, 29017 Fiorenzuola d'Arda, Italy (A.T., L.C.);Hubei Provincial Key Laboratory for Protection and Application of Special Plants in Wu Ling Area of China, Key Laboratory of State Ethnic Affairs Commission for Biologica Technology, College of Life Science, South-Central University for Nationalities, Wuhan 430074, China (X.X.); andParco Tecnologico Padano, Loc. Cascina Codazza, 26900 Lodi, Italy (L.R.) laura.rossini@unimi.it korff@mpipz.mpg.de.
authors.forename: Maria
authors.identifier: 0000-0002-6816-586X
authors.initials: M
authors.lastname: von Korff
mesh_terms: D002874:Chromosome Mapping; D032461:Chromosomes, Plant / Q000235:genetics; D035264:Flowers / Q000235:genetics; D018507:Gene Expression Regulation, Developmental; D018506:Gene Expression Regulation, Plant; D017343:Genes, Plant / Q000235:genetics*; D018745:Genome, Plant / Q000235:genetics; D005838:Genotype; D001467:Hordeum / Q000033:anatomy & histology / Q000235:genetics* / Q000254:growth & development; D015810:Linkage Disequilibrium; D017440:Photoperiod; D018515:Plant Leaves / Q000033:anatomy & histology / Q000235:genetics* / Q000254:growth & development; D020641:Polymorphism, Single Nucleotide; D040641:Quantitative Trait Loci / Q000235:genetics*; D020133:Reverse Transcriptase Polymerase Chain Reaction; D013997:Time Factors