title: Pore size regulates operating stomatal conductance, while stomatal densities drive the partitioning of conductance between leaf sides.
abstract: OBJECTIVE
  Leaf gas exchange is influenced by stomatal size, density, distribution between the leaf adaxial and abaxial sides, as well as by pore dimensions. This study aims to quantify which of these traits mainly underlie genetic differences in operating stomatal conductance (gs) and addresses possible links between anatomical traits and regulation of pore width.
  
  
  METHODS
  Stomatal responsiveness to desiccation, gs-related anatomical traits of each leaf side and estimated gs (based on these traits) were determined for 54 introgression lines (ILs) generated by introgressing segments of Solanum pennelli into the S. lycopersicum 'M82'. A quantitative trait locus (QTL) analysis for stomatal traits was also performed.
  
  
  RESULTS
  A wide genetic variation in stomatal responsiveness to desiccation was observed, a large part of which was explained by stomatal length. Operating gs ranged over a factor of five between ILs. The pore area per stomatal area varied 8-fold among ILs (2-16 %), and was the main determinant of differences in operating gs between ILs. Operating gs was primarily positioned on the abaxial surface (60-83 %), due to higher abaxial stomatal density and, secondarily, to larger abaxial pore area. An analysis revealed 64 QTLs for stomatal traits in the ILs, most of which were in the direction of S. pennellii.
  
  
  CONCLUSIONS
  The data indicate that operating and maximum gs of non-stressed leaves maintained under stable conditions deviate considerably (by 45-91 %), because stomatal size inadequately reflects operating pore area (R(2) = 0·46). Furthermore, it was found that variation between ILs in both stomatal sensitivity to desiccation and operating gs is associated with features of individual stoma. In contrast, genotypic variation in gs partitioning depends on the distribution of stomata between the leaf adaxial and abaxial epidermis.
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil dimitrios.fanourakis82@gmail.com.
authors.forename: Dimitrios
authors.initials: D
authors.lastname: Fanourakis
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Habtamu
authors.initials: H
authors.lastname: Giday
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Rubén
authors.initials: R
authors.lastname: Milla
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Roland
authors.initials: R
authors.lastname: Pieruschka
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Katrine H
authors.initials: KH
authors.lastname: Kjaer
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Marie
authors.initials: M
authors.lastname: Bolger
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Aleksandar
authors.initials: A
authors.lastname: Vasilevski
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Adriano
authors.initials: A
authors.lastname: Nunes-Nesi
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Fabio
authors.initials: F
authors.lastname: Fiorani
authors.affiliation: IBG-2: Plant Sciences, Institute for Bio- and Geosciences, Forschungszentrum Jülich, D-52425 Jülich, Germany, Aarhus University, Department of Food Science, Kirstinebjergvej 10, DK-5792 Årslev, Denmark, 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, Institute for Biology I, RWTH Aachen University, Aachen, Germany and Max Planck Partner Group at the Departamento de Biologia Vegetal, Universidade Federal de Viçosa, 36570-000 Viçosa, MG, Brazil.
authors.forename: Carl-Otto
authors.initials: CO
authors.lastname: Ottosen
mesh_terms: D003890:Desiccation; D014644:Genetic Variation; D006824:Hybridization, Genetic; D018551:Solanum lycopersicum / Q000033:anatomy & histology / Q000235:genetics / Q000502:physiology; D008954:Models, Biological; D010641:Phenotype; D018515:Plant Leaves / Q000033:anatomy & histology / Q000502:physiology*; D054046:Plant Stomata / Q000033:anatomy & histology / Q000502:physiology*; D032322:Solanum / Q000033:anatomy & histology / Q000235:genetics / Q000502:physiology*
keywords: Amphistomatous; QTL; S. pennellii; Solanum lycopersicum; leaf gas exchange; operating stomatal conductance; pore area; quantitative trait locus; stomatal responsiveness