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Silva Fennica 1926-1997
Acta Forestalia Fennica

Articles by Ilkka Leinonen

Category: Article

article id 5590, category Article
Heikki Hänninen, Seppo Kellomäki, Ilkka Leinonen, Tapani Repo. (1996). Overwintering and productivity of Scots pine in a changing climate. Silva Fennica vol. 30 no. 2–3 article id 5590.
Keywords: Pinus sylvestris; climate change; ecosystems; frost damage; phenology; forest ecology; gap-type model; model accuracy; model realism; overvintering
Abstract | View details | Full text in PDF | Author Info

The productivity of Scots pine (Pinus sylvestris L.) under changing climatic conditions in the southern part of Finland was studied by scenario analysis with a gap-type forest ecosystem model. Standard simulations with the model predicted an increased rate of growth and hence increased productivity as a result of climatic warming. The gap-type model was refined by introducing an overwintering sub-model describing the annual growth cycle, frost hardiness, and frost damage of the trees. Simulations with the refined gap-type model produced results conflicting with those of the standard simulation, i.e., drastically decreased productivity caused by mortality and growth-reducing damage due to premature dehardening in the changing climate. The overwintering sub-model was tested with frost hardiness data from Scots pine saplings growing at their natural site 1) under natural conditions and 2) under elevated temperature condition, both in open-top chambers. The model predicted the frost hardiness dynamics quite accurately for the natural conditions while underestimating the frost hardiness of the saplings for the elevated temperature conditions. These findings show that 1) the overwintering sub-model requires further development, and 2) the possible reduction of productivity caused by frost damage in a changing climate is less drastic than predicted in the scenario analysis. The results as a whole demonstrated the need to consider the overwintering of trees in scenario analysis carried out with ecosystem model for boreal conditions. More generally, the results revealed a problem that exists in scenario analysis with ecological models: the accuracy of a model in predicting the ecosystem functioning under present climatic condition does not guarantee the realism of the model, nor for this reason the accuracy for predicting the ecosystem functioning under changing climatic conditions. This finding calls for the continuous rigorous experimental testing of ecological models used for assessing the ecological implications of climatic change.

  • Hänninen, E-mail: hh@mm.unknown (email)
  • Kellomäki, E-mail: sk@mm.unknown
  • Leinonen, E-mail: il@mm.unknown
  • Repo, E-mail: tr@mm.unknown
article id 5583, category Article
Ilkka Leinonen, Heikki Hänninen, Tapani Repo. (1996). Testing of frost hardiness models for Pinus sylvestris in natural conditions and in elevated temperature. Silva Fennica vol. 30 no. 2–3 article id 5583.
Keywords: Pinus sylvestris; climate change; frost hardiness; Scots pine; temperature; dynamic models; photoperiod; climatic warming
Abstract | View details | Full text in PDF | Author Info

Two dynamic models predicting the development of frost hardiness of Finnish Scots pine (Pinus sylvestris L.) were tested with frost hardiness data obtained from trees growing in the natural conditions of Finland and from an experiment simulating the predicted climatic warming. The input variables were temperature in the first model, and temperature and night length in the second. The model parameters were fixed on the basis of previous independent studies. The results suggested that the model which included temperature and photoperiod as input variables was more accurate than the model using temperature as the only input variable to predict the development of frost hardiness in different environmental conditions. Further requirements for developing the frost hardiness models are discussed.

  • Leinonen, E-mail: il@mm.unknown (email)
  • Hänninen, E-mail: hh@mm.unknown
  • Repo, E-mail: tr@mm.unknown

Category: Research note

article id 534, category Research note
Ilkka Leinonen, Heikki Hänninen. (2002). Adaptation of the timing of bud burst of Norway spruce to temperate and boreal climates. Silva Fennica vol. 36 no. 3 article id 534.
Keywords: adaptation; modelling; frost damage; bud burst; phenology; stabilizing selection
Abstract | View details | Full text in PDF | Author Info
The adaptation of the annual cycle of development of boreal and temperate trees to climatic conditions has been seen as a result of stabilizing selection caused by two opposite driving forces of natural selection, i.e. the tolerance of unfavorable conditions during the frost exposed season (survival adaptation) and the effective use of growth resources during the growing season (capacity adaptation). In this study, two theories of the effects of climate on the adaptation of the timing of bud burst of trees were evaluated. This was done with computer simulations by applying a temperature sum model for predicting the timing of bud burst of different Norway spruce genotypes on the basis of air temperature data from various climatic conditions. High geographical variation in the temperature response of bud burst, typical for Norway spruce, was included in the theoretical analyses. The average timing of bud burst and the corresponding risk of occurrence of damaging frost during the susceptible period after bud burst were calculated for each genotype in each climate. Two contrasting theories of the stabilizing selection were evaluated, i.e. the overall adaptedness of each genotype was evaluated either 1) by assuming a fixed threshold for the risk of frost damage, or 2) by assuming a tradeoff between the risk of frost damage and the length of the growing season. The tradeoff assumption produced predictions of between provenance variation in bud burst which correspond more closely with empirical observations available in literature, compared to the fixed threshold assumption.
  • Leinonen, University of Oklahoma, Department of Botany and Microbiology, Norman, OK 73019, USA E-mail: (email)
  • Hänninen, University of Helsinki, Department of Ecology and Systematics, FIN-00014 Helsinki, Finland E-mail:
article id 608, category Research note
Jouni Partanen, Ilkka Leinonen, Tapani Repo. (2001). Effect of accumulated duration of the light period on bud burst in Norway spruce (Picea abies) of varying ages. Silva Fennica vol. 35 no. 1 article id 608.
Keywords: Norway spruce; phenology; growth initiation; photoperiod
Abstract | View details | Full text in PDF | Author Info
One-year-old seedlings (two sowing times), two-year-old seedlings and 14- and 18-year-old cuttings of Norway spruce (Picea abies (L.) Karst.) were exposed to shortening photoperiod (initially 16 h), lengthening photoperiod (initially 6 h) and constant short photoperiod (6 h) treatments with uniform temperature conditions in growth chambers. The timing of bud burst was examined. In all plants, shortening photoperiod treatment seemed to promote bud burst compared with other treatments. This effect was clearest in the oldest material. The results suggest that, in addition to temperature sum, the accumulated duration of the light period may promote bud burst of Norway spruce.
  • Partanen, Finnish Forest Research Institute, Punkaharju Research Station, Finlandiantie 18, FIN-58450 Punkaharju, Finland E-mail: (email)
  • Leinonen, University of Joensuu, Faculty of Forestry, P.O. Box 111, FIN-80101 Joensuu, Finland E-mail:
  • Repo, University of Joensuu, Faculty of Forestry, P.O. Box 111, FIN-80101 Joensuu, Finland E-mail:

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