title: Leaf dynamics in growth and reproduction of Xanthium canadense as influenced by stand density.
abstract: OBJECTIVE
  Leaf longevity is controlled by the light gradient in the canopy and also by the nitrogen (N) sink strength in the plant. Stand density may influence leaf dynamics through its effects on light gradient and on plant growth and reproduction. This study tests the hypothesis that the control by the light gradient is manifested more in the vegetative period, whereas the opposite is true when the plant becomes reproductive and develops a strong N sink.
  
  
  METHODS
  Stands of Xanthium canadense were established at two densities. Emergence, growth and death of every leaf on the main stem and branches, and plant growth and N uptake were determined from germination to full senescence. Mean residence time and dry mass productivity were calculated per leaf number, leaf area, leaf mass and leaf N (collectively termed 'leaf variables') in order to analyse leaf dynamics and its effect on plant growth.
  
  
  RESULTS
  Branching and reproductive activities were higher at low than at high density. Overall there was no significant difference in mean residence time of leaf variables between the two stands. However, early leaf cohorts on the main stem had a longer retention time at low density, whereas later cohorts had a longer retention time at high density. Branch leaves emerged earlier and tended to live longer at low than at high density. Leaf efficiencies, defined as carbon export per unit investment of leaf variables, were higher at low density in all leaf variables except for leaf number.
  
  
  CONCLUSIONS
  In the vegetative phase of plant growth, the light gradient strongly controls leaf longevity, whereas later the effects of branching and reproductive activities become stronger and over-rule the effect of light environment. As leaf N supports photosynthesis and also works as an N source for plant development, N use is pivotal in linking leaf dynamics with plant growth and reproduction.
authors.affiliation: Department of International Agricultural Development, Tokyo University of Agriculture, Tokyo 156-8502, Japan.
authors.forename: Takahiro
authors.initials: T
authors.lastname: Ogawa
authors.affiliation: Department of International Agricultural Development, Tokyo University of Agriculture, Tokyo 156-8502, Japan.
authors.forename: Shimpei
authors.initials: S
authors.lastname: Oikawa
authors.affiliation: Department of International Agricultural Development, Tokyo University of Agriculture, Tokyo 156-8502, Japan tadakihirose@gmail.com.
authors.forename: Tadaki
authors.initials: T
authors.lastname: Hirose
mesh_terms: D008027:Light; D009584:Nitrogen / Q000378:metabolism*; D010788:Photosynthesis; D018515:Plant Leaves / Q000254:growth & development / Q000502:physiology*; D011156:Population Density; D031210:Xanthium / Q000254:growth & development / Q000502:physiology*
chemical_list: D009584:Nitrogen
keywords: Asteraceae; Leaf dynamics; Xanthium canadense; cocklebur; leaf longevity; leaf productivity; mean residence time; nitrogen resorption; nitrogen use efficiency; stand density