title: Overwintering leaves of a forest-floor fern, Dryopteris crassirhizoma (Dryopteridaceae): a small contribution to the resource storage and photosynthetic carbon gain.
abstract: OBJECTIVE
  Dryopteris crassirhizoma is a semi-evergreen fern growing on the floor of deciduous forests. The present study aimed to clarify the photosynthetic and storage functions of overwintering leaves in this species.
  
  
  METHODS
  A 2-year experiment with defoliation and shading of overwintering leaves was conducted. Photosynthetic light response was measured in early spring (for overwintering leaves) and summer (for current-year leaves).
  
  
  RESULTS
  No nitrogen limitation of growth was detected in plants subjected to defoliation. The number of leaves, their size, reproductive activity (production of sori) and total leaf mass were not affected by the treatment. The defoliation of overwintering leaves significantly reduced the bulk density of rhizomes and the root weight. The carbohydrates consumed by the rhizomes were assumed to be translocated for leaf production. Photosynthetic products of overwintering leaves were estimated to be small.
  
  
  CONCLUSIONS
  Overwintering leaves served very little as nutrient-storage and photosynthetic organs. They partly functioned as a carbon-storage organ but by contrast to previous studies, their physiological contribution to growth was found to be modest, probably because this species has a large rhizome system. The small contribution of overwintering leaves during the short-term period of this study may be explained by the significant storage ability of rhizomes in this long-living species. Other ecological functions of overwintering leaves, such as suppression of neighbouring plants in spring, are suggested.
authors.affiliation: Graduate School of Environmental Earth Science, Hokkaido University, Sapporo 060-0810, Japan.
authors.forename: Tomokazu
authors.initials: T
authors.lastname: Tani
authors.forename: Gaku
authors.initials: G
authors.lastname: Kudo
mesh_terms: D002244:Carbon / Q000378:metabolism*; D032492:Dryopteris / Q000502:physiology*; D004777:Environment; D010788:Photosynthesis / Q000502:physiology*; D018515:Plant Leaves / Q000502:physiology*; D012621:Seasons
chemical_list: D002244:Carbon