title: Changes in leaf trichomes and epicuticular flavonoids during leaf development in three birch taxa.
abstract: OBJECTIVE
  Changes in number of trichomes and in composition and concentrations of their exudates throughout leaf development may have important consequences for plant adaptation to abiotic and biotic factors. In the present study, seasonal changes in leaf trichomes and epicuticular flavonoid aglycones in three Finnish birch taxa (Betula pendula, B. pubescens ssp. pubescens, and B. pubescens ssp. czerepanovii) were followed.
  
  
  METHODS
  Trichome number and ultrastructure were studied by means of light, scanning and transmission electron microscopy, while flavonoid aglycones in ethanolic leaf surface extracts were analysed by high-pressure liquid chromatography.
  
  
  RESULTS
  Density of both glandular and non-glandular trichomes decreased drastically with leaf expansion while the total number of trichomes per leaf remained constant, indicating that the final number of trichomes is established early in leaf development. Cells of glandular trichomes differentiate before those of the epidermis and produce secreted material only during the relatively short period (around 1-2 weeks) of leaf unfolding and expansion. In fully expanded leaves, glandular trichomes appeared to be at the post-secretory phase and function mainly as storage organs; they contained lipid droplets and osmiophilic material (probably phenolics). Concentrations (mg g(-1) d. wt) of surface flavonoids decreased with leaf age in all taxa. However, the changes in total amount ( microg per leaf) of flavonoids during leaf development were taxon-specific: no changes in B. pubescens ssp. czerepanovii, increase in B. pendula and in B. pubescens ssp. pubescens followed by the decline in the latter taxon. Concentrations of most of the individual leaf surface flavonoids correlated positively with the density of glandular trichomes within species, suggesting the participation of glandular trichomes in production of surface flavonoids.
  
  
  CONCLUSIONS
  Rapid decline in the density of leaf trichomes and in the concentrations of flavonoid aglycones with leaf age suggests that the functional role of trichomes is likely to be most important at the early stages of birch leaf development.
authors.affiliation: Section of Ecology, Department of Biology, University of Turku, Finland. elekos@utu.fi
authors.forename: Elena
authors.initials: E
authors.lastname: Valkama
authors.forename: Juha-Pekka
authors.initials: JP
authors.lastname: Salminen
authors.forename: Julia
authors.initials: J
authors.lastname: Koricheva
authors.forename: Kalevi
authors.initials: K
authors.lastname: Pihlaja
mesh_terms: D029662:Betula / Q000145:classification / Q000254:growth & development* / Q000648:ultrastructure; D022081:Cell Surface Extensions / Q000502:physiology* / Q000648:ultrastructure; D005419:Flavonoids / Q000378:metabolism*; D008855:Microscopy, Electron, Scanning; D019441:Plant Epidermis / Q000254:growth & development* / Q000378:metabolism; D018515:Plant Leaves / Q000254:growth & development* / Q000378:metabolism / Q000648:ultrastructure
chemical_list: D005419:Flavonoids