title: Warming differentially influences the effects of drought on stoichiometry and metabolomics in shoots and roots.
abstract: Plants in natural environments are increasingly being subjected to a combination of abiotic stresses, such as drought and warming, in many regions. The effects of each stress and the combination of stresses on the functioning of shoots and roots have been studied extensively, but little is known about the simultaneous metabolome responses of the different organs of the plant to different stresses acting at once. We studied the shift in metabolism and elemental composition of shoots and roots of two perennial grasses, Holcus lanatus and Alopecurus pratensis, in response to simultaneous drought and warming. These species responded differently to individual and simultaneous stresses. These responses were even opposite in roots and shoots. In plants exposed to simultaneous drought and warming, terpenes, catechin and indole acetic acid accumulated in shoots, whereas amino acids, quinic acid, nitrogenous bases, the osmoprotectants choline and glycine betaine, and elements involved in growth (nitrogen, phosphorus and potassium) accumulated in roots. Under drought, warming further increased the allocation of primary metabolic activity to roots and changed the composition of secondary metabolites in shoots. These results highlight the plasticity of plant metabolomes and stoichiometry, and the different complementary responses of shoots and roots to complex environmental conditions.
authors.affiliation: Global Ecology Unit CREAF-CSIC-UAB, CSIC, Cerdanyola del vallès, Catalonia, 08193, Spain.|CREAF, Cerdanyola del vallès, Catalonia, 08193, Spain.|Service of Nuclear Magnetic Resonance and Chemistry Department, Faculty of Sciences and Biosciences, Universitat Autònoma de Barcelona, Bellaterra, Catalonia, 08913, Spain.
authors.forename: Albert
authors.initials: A
authors.lastname: Gargallo-Garriga
authors.affiliation: Global Ecology Unit CREAF-CSIC-UAB, CSIC, Cerdanyola del vallès, Catalonia, 08193, Spain.|CREAF, Cerdanyola del vallès, Catalonia, 08193, Spain.
authors.forename: Jordi
authors.initials: J
authors.lastname: Sardans
authors.affiliation: Service of Nuclear Magnetic Resonance and Chemistry Department, Faculty of Sciences and Biosciences, Universitat Autònoma de Barcelona, Bellaterra, Catalonia, 08913, Spain.
authors.forename: Míriam
authors.initials: M
authors.lastname: Pérez-Trujillo
authors.affiliation: Global Change Research Centre, Academy of Sciences of the Czech Republic, Belidla 4a, CZ-60300, Brno, Czech Republic.
authors.forename: Michal
authors.initials: M
authors.lastname: Oravec
authors.affiliation: Global Change Research Centre, Academy of Sciences of the Czech Republic, Belidla 4a, CZ-60300, Brno, Czech Republic.
authors.forename: Otmar
authors.initials: O
authors.lastname: Urban
authors.affiliation: Disturbance Ecology and Vegetation Dynamics, University of Bayreuth, Bayreuth, D-95440, Germany.
authors.forename: Anke
authors.initials: A
authors.lastname: Jentsch
authors.affiliation: Department of Biogeography, University of Bayreuth, Bayreuth, D-95440, Germany.
authors.forename: Juergen
authors.initials: J
authors.lastname: Kreyling
authors.affiliation: Department of Biogeography, University of Bayreuth, Bayreuth, D-95440, Germany.
authors.forename: Carl
authors.initials: C
authors.lastname: Beierkuhnlein
authors.affiliation: Service of Nuclear Magnetic Resonance and Chemistry Department, Faculty of Sciences and Biosciences, Universitat Autònoma de Barcelona, Bellaterra, Catalonia, 08913, Spain.
authors.forename: Teodor
authors.initials: T
authors.lastname: Parella
authors.affiliation: Global Ecology Unit CREAF-CSIC-UAB, CSIC, Cerdanyola del vallès, Catalonia, 08193, Spain.|CREAF, Cerdanyola del vallès, Catalonia, 08193, Spain.
authors.forename: Josep
authors.initials: J
authors.lastname: Peñuelas
mesh_terms: D016002:Discriminant Analysis; D055864:Droughts; D004602:Elements; D031752:Holcus / Q000378:metabolism*; D006358:Hot Temperature; D016018:Least-Squares Analysis; D055442:Metabolome; D055432:Metabolomics; D018517:Plant Roots / Q000378:metabolism*; D018520:Plant Shoots / Q000378:metabolism*; D006109:Poaceae / Q000378:metabolism*; D025341:Principal Component Analysis; D012621:Seasons; D013045:Species Specificity
chemical_list: D004602:Elements
keywords: HPLC-MS; climate change; drought; metabolomics; nitrogen : phosphorus (N : P); nuclear magnetic resonance (NMR); stoichiometry; warming