title: Shifts and disruptions in resource-use trait syndromes during the evolution of herbaceous crops.
abstract: Trait-based ecology predicts that evolution in high-resource agricultural environments should select for suites of traits that enable fast resource acquisition and rapid canopy closure. However, crop breeding targets specific agronomic attributes rather than broad trait syndromes. Breeding for specific traits, together with evolution in high-resource environments, might lead to reduced phenotypic integration, according to predictions from the ecological literature. We provide the first comprehensive test of these hypotheses, based on a trait-screening programme of 30 herbaceous crops and their wild progenitors. During crop evolution plants became larger, which enabled them to compete more effectively for light, but they had poorly integrated phenotypes. In a subset of six herbaceous crop species investigated in greater depth, competitiveness for light increased during early plant domestication, whereas diminished phenotypic integration occurred later during crop improvement. Mass-specific leaf and root traits relevant to resource-use strategies (e.g. specific leaf area or tissue density of fine roots) changed during crop evolution, but in diverse and contrasting directions and magnitudes, depending on the crop species. Reductions in phenotypic integration and overinvestment in traits involved in competition for light may affect the chances of upgrading modern herbaceous crops to face current climatic and food security challenges.
authors.affiliation: Departamento de Biología y Geología, Área de Biodiversidad y Conservación, Escuela Superior de Ciencias Experimentales y Tecnología, Universidad Rey Juan Carlos, c/Tulipán s/n, Móstoles 28933, Spain ruben.milla@gmail.com.
authors.forename: Rubén
authors.initials: R
authors.lastname: Milla
authors.affiliation: Departamento de Biología y Geología, Área de Biodiversidad y Conservación, Escuela Superior de Ciencias Experimentales y Tecnología, Universidad Rey Juan Carlos, c/Tulipán s/n, Móstoles 28933, Spain.
authors.forename: Javier
authors.initials: J
authors.lastname: Morente-López
authors.affiliation: Departamento de Biología y Geología, Área de Biodiversidad y Conservación, Escuela Superior de Ciencias Experimentales y Tecnología, Universidad Rey Juan Carlos, c/Tulipán s/n, Móstoles 28933, Spain.
authors.forename: J Miguel
authors.initials: JM
authors.lastname: Alonso-Rodrigo
authors.affiliation: Departamento de Biología y Geología, Área de Biodiversidad y Conservación, Escuela Superior de Ciencias Experimentales y Tecnología, Universidad Rey Juan Carlos, c/Tulipán s/n, Móstoles 28933, Spain.
authors.forename: Nieves
authors.initials: N
authors.lastname: Martín-Robles
authors.affiliation: Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, AK 99775, USA.
authors.forename: F Stuart
authors.initials: FS
authors.lastname: Chapin
mesh_terms: D005075:Biological Evolution; D001947:Breeding; D018556:Crops, Agricultural / Q000235:genetics / Q000502:physiology*; D010641:Phenotype; D018515:Plant Leaves / Q000502:physiology; D018517:Plant Roots / Q000502:physiology
keywords: breeding; competition; domestication; phenotypic integration; plant ecological strategies; structural equation models