Cold adaptation drives variability in needle structure and anatomy in Pinus sylvestris L. along a 1,900 km temperate-boreal transect

Research output: Contribution to journalArticle


Occupancy of cold habitats by evergreen species requires structural modification of photosynthetic organs for stress resistance and longevity. Such modifications have been described at interspecific level, while intraspecific variation has been underexplored. To identify structural and anatomical traits that may be adaptive in cold environments, we studied intraspecific variability of needles of Scots pine (Pinus sylvestris L.), a wide-ranging tree, along a 1,900 km temperate-boreal transect in Europe. Needles from 20 sites representing mean minimum winter temperatures between -4.0 and -19.9°C and mean annual temperatures between 8.3 and -1.7°C were sampled for measurements of leaf mass per area (LMA, g/m2), leaf density (LD, g/cm3) and 30 other morpho-anatomical traits. Needles from cold sites lived longer, were shorter, showed higher LMA and LD, had narrower and more collapse-resistant tracheids, thicker epidermal cells with thicker cell walls and wider resin ducts occupying larger fraction of needle volume in comparison to needles from warmer sites. Along the steep climatic gradient, needles presented a coordinated phenotypic spectrum of external and internal traits that are largely interpretable in functional, adaptive terms. This intraspecific pattern of covarying traits provides insight into the adaptive syndrome associated with stress tolerance and extended needle longevity under cold conditions of high latitudes. A plain language summary is available for this article.


  • Artur Jankowski
  • Tomasz P. Wyka
  • Roma Zytkowiak
  • Bengt Nihlgård
  • Peter B. Reich
  • Jacek Oleksyn
External organisations
  • University of Minnesota system
  • Adam Mickiewicz University in Poznań
  • Institute of Dendrology, PAS
  • Western Sydney University
Research areas and keywords

Subject classification (UKÄ) – MANDATORY

  • Evolutionary Biology


  • Anatomical variation, Foliar tracheids, Leaf economic spectrum, Leaf longevity, Resin ducts
Original languageEnglish
Pages (from-to)2212-2223
JournalFunctional Ecology
Issue number12
Early online date2017 Sep 11
Publication statusPublished - 2017 Dec
Publication categoryResearch