title: Chemical and physical defence in early and late leaves in three heterophyllous birch species native to northern Japan.
abstract: OBJECTIVE
  Betula ermanii, B. maximowicziana and B. platyphylla var. japonica have heterophyllous leaves (i.e. early leaves and late leaves) and are typical pioneer species in northern Japan. Chemical and physical defences against herbivores in early and late leaves of these species were studied.
  
  
  METHODS
  Two-year-old seedlings were grown under full sunlight in a single growing season. Three-week-old leaves of each seedling were harvested three times (May, July and October). Total phenolics and condensed tannin content were determined for chemical defence and leaf toughness and trichome density were assessed for physical defence. Defoliation of early leaves in May was also performed to study the contribution of early leaves to subsequent growth.
  
  
  RESULTS
  Chemical and physical defences were greater in early than late leaves in B. platyphylla and B. ermanii, whereas the reverse was true in B. maximowicziana. In contrast to its weak chemical defences, the trichome density in B. maximowicziana was very high. In B. platyphylla and B. ermanii, the relative growth rates (RGR) were greater early in the growing season. Negative effects on growth of removal of early leaves were significant only in B. platyphylla.
  
  
  CONCLUSIONS
  B. platyphylla and B. ermanii invest in defence in early rather than late leaves, since early leaves are crucial to subsequent growth. In contrast, B. maximowicziana more strongly defends its late leaves, since its RGR is maintained at the same level throughout the growing season.
authors.affiliation: Graduate School of Agriculture, Hokkaido University, Sapporo 060-8589, Japan.
authors.forename: Sawako
authors.initials: S
authors.lastname: Matsuki
authors.forename: Yuzou
authors.initials: Y
authors.lastname: Sano
authors.forename: Takayoshi
authors.initials: T
authors.lastname: Koike
mesh_terms: D029662:Betula / Q000502:physiology*; D005843:Geography; D007564:Japan; D018515:Plant Leaves / Q000166:cytology / Q000254:growth & development / Q000502:physiology*; D012621:Seasons; D013472:Sunlight