title: Abaxial growth and steric constraints guide leaf folding and shape in Acer pseudoplatanus.
abstract: OBJECTIVE
  How leaf shape is regulated is a long-standing question in botany. For diverse groups of dicotyledon species, lamina folding along the veins and geometry of the space available for the primordia can explain the palmate leaf morphology. Dubbed the kirigami theory, this hypothesis of fold-dependent leaf shape regulation has remained largely theoretical. Using Acer pseudoplatanus, we investigated the mechanisms behind the two key processes of kirigami leaf development.
  
  
  METHODS
  Cytological examination and quantitative analyses were used to examine the course of the vein-dependent lamina folding. Surgical ablation and tissue culturing were employed to test the effects of physical constraints on primordia growth. The final morphology of leaves growing without steric constraints were predicted mathematically.
  
  
  RESULTS
  The cytological examination showed that the lamina's abaxial side along the veins grows substantially more than the adaxial side. The abaxial hypergrowth along the veins and the lamina extension correlated with the lamina folding. When a primordium was released from the physical constraints imposed by the other primordia, it rapidly grew into the newly available space, while maintaining the curvature inward. The morphology of such a leaf was predicted to lack symmetry in the lobe shapes.
  
  
  CONCLUSIONS
  The enhanced growth on the abaxial side of the lamina along the veins is likely to drive lamina folding. The surgical ablation provided clear support for the space-filling nature of leaf growth; thus, steric constraints play a role in determination of the shapes of folded leaves and probably also of the final leaf morphology.
authors.affiliation: Laboratoire MSC, UMR CNRS/Université Paris-Diderot, France.
authors.forename: Etienne
authors.initials: E
authors.lastname: Couturier
authors.forename: Nicole
authors.initials: N
authors.lastname: Brunel
authors.forename: Stéphane
authors.initials: S
authors.lastname: Douady
authors.forename: Naomi
authors.initials: N
authors.lastname: Nakayama
mesh_terms: D031002:Acer / Q000166:cytology* / Q000254:growth & development*; D001696:Biomechanical Phenomena; D008962:Models, Theoretical; D018515:Plant Leaves / Q000166:cytology* / Q000254:growth & development*; D018520:Plant Shoots / Q000166:cytology / Q000254:growth & development