[1] Nat Resources Inst Finland LUKE, Prod Syst, Joensuu 80100, North Karelia, Finland
[2] Univ Eastern Finland, Sch Forest Sci, Joensuu 80100, North Karelia, Finland
[3] Nat Resources Inst Finland LUKE, Nat Resources, Helsinki 00790, Finland
[4] Univ Pretoria, Forestry & Agr Biotechnol Inst FABI, Dept Biochem Genet & Microbiol, ZA-0028 Pretoria, South Africa