Articles by Eero Nikinmaa

The effects of two alternative formulations of sapwood senescence on the behaviour of model LIGNUM (with parameter values adjusted for Scots pine (Pinus sylvestris L.) growing southern Finland) were studied. The two alternatives were autonomous sapwood senescence assuming a maximum age for the tree ring, and sapwood senescence that is controlled by the mortality of foliage. For the latter alternative two hypothetical further mechanisms were stipulated. All the formulations were implemented in LIGNUM. Simulations were made with all model variants for fertile and poor soil conditions using high, normal and low rates of foliage mortality. The simulation results were compared against of a data set consisting of 11 open grown Scots pine trees from southern Finland. Observations of heartwood proportion were used in this study. They show that heartwood starts to increase in trees from age of approximately 20 years onwards. The simulation results showed no differences between fertile and poor soil conditions as regards heartwood formation. Of the variants of foliage-controlled sapwood senescence the one where death of sapwood in a tree segment induces sapwood senescence in the tree parts below only slightly was the best. This and the autonomous sapwood senescence corresponded equally well to the observations. In order to make more refined conclusions additional data and simulations are necessary.

• Sievänen, E-mail: rs@mm.unknown
• Nikinmaa, E-mail: en@mm.unknown
• Perttunen, E-mail: jp@mm.unknown

A theoretical framework to analyse the growth of Scots pine (Pinus sylvestris L.) is presented. Material exchange processes and internal processes that transport, transform and consume materials are identified as the components of growth. Hierarchical system is lined out. Momentary uptake of material at a single exchange site depends on the environmental condition next to the exchange site, the internal state of the biochemical system of the plant and the structure of the plant. The internal state depends on the exchange flows over period of time and the structural growth depends on the internal state. The response of these processes to the fluxes is controlled by the genetic composition of the plant.

The theoretical framework is formulated into a mathematical model. A concept of balanced internal state was applied to describe the poorly known internal processes. Internal substrate concentrations were assumed to remain constant but tissue-specific. A linear relationship between the quantity of foliage and wood cross-sectional area was assumed to describe balanced formation of structure. The exchange processes were thus described as a function of external conditions. The stand level interactions were derived from shading and effects of root density on nutrient uptake.

The approach was tested at different levels of hierarchy. Field measurements indicated that the hypothesis of the linear relationship described well the regularities between foliage and sapwood of a tree within a stand when measured at functionally corresponding height. There was considerable variation in the observed regularities in the range of geographic occurrence of Scots pine. Model simulations gave a realistic description of stand development in Southern Finland. The same model was also able to describe growth differences in Lapland after considering the effect of growing season length in the parameter values. Simulations to South Russia indicate stronger deviation from the observed patterns.

The simulations suggest interesting features of stand development. They indicate strong variability in the distribution of carbohydrates between tree parts during stand development. Internal circulation of nutrients and the reuse of the same transport structure by various needle generations had a strong influence on the simulation results.

The PDF includes a summary in Finnish.

• Nikinmaa, E-mail: en@mm.unknown

Despite the numerous studies on year-to-year variation of tree growth, the physiological mechanisms controlling annual variation in growth are still not understood in detail. We studied the applicability of data-driven approach i.e. different regression models in analysing high-dimensional data set including continuous and comprehensive measurements over meteorology, ecosystem-scale water and carbon fluxes and the annual variation in the growth of app. 50-year-old Scots pine stand in southern Finland. Even though our dataset covered only 16 years, it is the most extensive collection of interactions between a Scots pine ecosystem and atmosphere. The analysis revealed that height growth was favoured by high water potential of the tree and carbon gain during the bud forming period and high water potential during the elongation period. Diameter growth seemed to be favoured by a winter with high precipitation and deep snow cover and a spring with high carbon gain. The obtained models had low generalization performance and they would require more evaluation and iterative validation to achieve credibility perhaps as a mixture of data-driven and first principle modeling approaches.

• Kulmala, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: liisa.kulmala@helsinki.fi
• Žliobaitė, Aalto University, Department of Computer Science and Helsinki Institute for Information Technology, P.O. Box 11000, FI-00076 Aalto, Finland; University of Helsinki, Department of Geosciences and Geography, P.O. Box 64, FI-00014 University of Helsinki, Finland E-mail: zliobaite@gmail.com
• Nikinmaa, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: eero.nikinmaa@helsinki.fi
• Nöjd, Natural Resources Institute Finland (Luke), Bio-based business and industry, Tietotie 2, FI-02150 Espoo, Finland E-mail: pekka.nojd@luke.fi
• Kolari, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland; University of Helsinki, Department of Physics, P.O. Box 64, FI-00014 University of Helsinki, Finland E-mail: pasi.kolari@helsinki.fi
• Kabiri Koupaei, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: kourosh.kabiri@helsinki.fi
• Hollmén, Aalto University, Department of Computer Science and Helsinki Institute for Information Technology, P.O. Box 11000, FI-00076 Aalto, Finland; University of Helsinki, Department of Geosciences and Geography, P.O. Box 64, FI-00014 University of Helsinki, Finland E-mail: jaakko.hollmen@aalto.fi
• Mäkinen, Natural Resources Institute Finland (Luke), Bio-based business and industry, Tietotie 2, FI-02150 Espoo, Finland E-mail: harri.makinen@luke.fi

• Kokkila, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FIN-00014 Helsingin yliopisto, Finland E-mail: tero.kokkila@helsinki.fi
• Mäkelä, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FIN-00014 Helsingin yliopisto, Finland E-mail: am@nn.fi
• Nikinmaa, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FIN-00014 Helsingin yliopisto, Finland E-mail: en@nn.fi