Current issue: 57(2)
Under compilation: 57(3)
The rule-based formal language of "stochastic sensitive growth grammars" was designed to describe algorithmically the changing morphology of forest trees during their lifetime under the impact of endogenous and exogenous factors, and to generate 3-D simulations of tree structures in a systematic manner. The description in the form of grammars allows the precise specification of structural models with functional components. These grammars (extended L-systems) can be interpreted by the software GROGRA (Growth grammar interpreter) yielding time series of attributed 3-D structures representing plants. With some recent extensions of the growth-grammar language (sensitive functions, local variables) it is possible to model environmental control of shoot growth and some simple allocation strategies, and to obtain typical competition effects in tree stands qualitatively in the model.
A metabolic model of height growth and site index is derived from a parametrization of the annual carbon balance of a tree. The parametrization is based on pipe-model theory. Four principal variants of the height-growth model correspond to four combinations of assumptions regarding carbon allocation: (a) the apical shoot is autonomous or (b) it is not; and (A) the specific rate of elongation of a shoot equals that of a woody root or (B) it does not. The bB model is the most general as it includes the aA, bA, and aB models as special cases. If the physiological parameters are constant, then the aA model reduces to the form of the Mitscherlich model and the bA model to the form of a Bertalanffy model. Responses of height growth to year-to-year variation in atmospheric conditions are rendered through adjustments of a subset of the model's parameters, namely, the specific rate of production of carbon substrate and three specific rates of maintenance respiration. As an example, the effect of the increasing atmospheric concentration of CO2 on the time-course of tree height of loblolly pine (Pinus taeda) is projected over 50-year span from 1986. Site index is predicted to increase and, more importantly, the shape of the site-index curve is predicted to change.
Process-based tree growth models are recognized to be flexible tools which are valuable for investigating tree growth in relation to changing environment or silvicultural treatments. In the context of forestry, we address two key modelling problems: allocation of growth which determines total wood production, and distribution of wood along the stem which determines stem form and wood quality. Growth allocation and distribution are the outcome of carbon translocation, which may be described by the Munch theory. We propose a simpler gradient process to describe the carbon distribution in the phloem of conifers. This model is a reformulation of a carbon diffusion-like process proposed by Thornley in 1972. By taking into account the continuity of the cambium along the stem, we obtain a one-dimensional reaction-diffusion model which describes both growth allocation between foliage, stem and roots, and growth distribution along the stem. Distribution of wood along the stem is then regarded as an allocation process at a smaller scale. A preliminary sensitivity analysis is presented. The model predicts a strong relationship between morphology and foliage-root allocation. It also suggests how empirical data, such as stem analysis, could be used to calibrate and validate allocation rules in process-based growth models.
Change in dry matter partitioning, 14C-incorporation, and sink 14C-activity of 1.5-year-old Scots pine (Pinus sylvestris L.) seedlings grown in growth chamber conditions were studied during a 91-day experiment. On five sampling dates, seedlings were labelled with 14CO2 and whole-plant allocation patterns were determined. Intensively growing shoots modified the dry matter partitioning: during shoot growth the proportion of roots decreased but after that it increased. Based on their large proportion of dry matter, the needles (excluding current needles) were the strongest sink of carbon containing 40% of the incorporated 14C. Despite their small initial sink size, the elongating shoots (current main shoot + current branch) and their needles were the second strongest sink (30–40% of the total 14C) which reflects their high physiological activity. The proportion of 14C in the current year’s main shoot increased during shoot growth but decreased as the growth began to decline after 70 days. 10–20% of the total assimilated 14C was translocated to the roots. Laterals above 2nd order were the strongest sink in the root system, containing twice as much 14C as the other roots together. Alternation between shoot and root growth can be seen clearly: carbon allocation to roots was relatively high before and after the period of intensive shoot growth. Changes in root sink strength resulted primarily from changes in root sink activity rather than sink size.
Crown characteristics and the distribution of three years’ (1986–88) biomass production of 20 pendulous Norway spruce (Picea abies f. pendula (Lawson) Sylvén) trees with heritable narrow crown and 15 normal-growned spruces (Picea abies (L.) H. Karst.) were studied in a 19-year-old mixed stand.
The form of the crown is conical in normal-crowned trees, columnar and narrow in pendulous trees. The partitioning of aboveground biomass to stems during the studied 3-year period was significantly higher in pendulous (0.281) than in normal-crowned trees (0.255) and also the ratio between growth of stemwood and growth of needle biomass during three years was higher in pendulous trees (0.67 g g-1) than in normal-crowned trees (0.52 g g-1). The needle biomass was distributed higher in the crown in pendula than in normal-crowned trees and they had a higher needle biomass/branchwood biomass ratio than normal trees. The difference in harvest increment between the two crown types are mostly due to the significantly lower branchwood biomass values in pendulous than in normal-crowned trees. The higher needle ’efficiency’ in pendulous trees is probably connected with high partitioning of needle biomass to the upper part of the crown in pendulous trees.
The PDF includes an abstract in Finnish.
A model was constructed, the aim of which was to predict growth under conditions where air pollutants are present. The model is based on photosynthesis and on the allocation of photosynthetic products for growth. It is assumed that air pollutants released during energy production mainly affect photosynthesis in two ways: 1) directly by injuring the photosynthetic mechanism, and 2) indirectly by leaching nutrients. The two ways were studied empirically in order to identify a sub-model for the photosynthesis of a plant exposed to air pollutants.
The stand model will be applied to two purposes. The present stage of forests in Finland is compared with the simulated state based on the assumption that no pollutants are present. In addition, the decrease in forest yield under different conditions derived from predictions about long-range pollutant transport in Europe is analysed.
The proportions of needle and wood in current-year shoots in crown systems of young Scots pine (Pinus sylvestris L.) trees was studied. The proportion of needles out of the total shoot biomass increased according to the increasing number of the whorl counted from the apex. In the lower part of the crown system the needle biomass of newly-formed shoots was 50–60 fold compared to that of wood and bark biomass. In the upper part of the crown system the same ratio was 1–2. The variation in ratio between needle and wood biomass was whorl-specific and independent of tree class. The magnitude of the ratio was not related to the position of the tree in the stand nor to the prevailing light conditions within the state.
The PDF includes a summary in Finnish.
One forest drainage undertaking in Finland often consists of woodlots belonging to several owners, and over hundred owners may be involved. In the present paper a method for allocation the costs to different owners in a joint drainage undertaking is worked out. The problem has been emphasised by the new Waterways Law, which enables also such drainage projects to be undertaken to which some of the land owners oppose. In those cased the costs must be allocated according to the benefit driven by each owner from the project.
The method attempts to assess the benefits to be driven from the forest drainage, those costs of the drainage that are joint and thus subjected to allocation, and what is the area affected by drainage as used as a basis for cost allocation.
The joined costs are apportioned in the following manner. The area of peatland adjusted to differences in the benefit obtained by drainage is ascertained by the land holder by multiplying the index number by the corresponding areas. In the case of cultivated agricultural land, also an index showing the need for drainage is used in computing the adjusted area. Each topographic unit in the map is provided with a notation of its apportionment area. Joined costs are allocated to different land owners in relation to their adjusted land areas.
The PDF includes a summary in English.
The accompanying collective research report is the result of the research project in 1986–90 between the Finnish Academy and the former Soviet Academy of Sciences. The project was organized around common field work in Finland and in the former Soviet Union, and theoretical analyses of tree growth determining processes. Based on theoretical analyses, dynamic stand growth models were made and their parameters were determined utilizing the field results.
Annual cycle effects the tree growth. Our theoretical approach was based on adaptation to local climate conditions from Lapland to South Russia. The initiation of growth was described as a simple low and high temperature accumulation driven model. The model was linked with long-term temperature data.
Analysis of field measurements of CO2 exchange showed that irradiance is the dominating factor causing variation in photosynthetic rate in natural conditions during summer. The penetration of irradiance into Scots pine (Pinus sylvestris L.) canopy is a complicated phenomenon. A moderately simple but balanced forest radiation regime sub-model was constructed.
The common field excursions in different geographical regions resulted in a lot of experimental data of regularities of woody structure. The water transport seems to be a good common factor to analyse these properties of tree structure. The produced theoretical and experimental material was utilized in the development of stand growth model that describes the growth and development of Scots pine stands in Finland and the former Soviet Union. The core of the model is carbon and nutrient balances. This means that carbon obtained in photosynthesis is consumed or growth and maintenance and nutrients are taken according to the metabolic need. Since the fundamental metabolic processes are the same in all locations, the same growth model structure can be applied in the large range of Scots pine. The model enables the analysis of geographical differences in the growth of Scots pine. The present approach enables utilization of structural and functional knowledge gained in places of intensive research, in the analysis of growth and development of any stand.
The aim of this study was to investigate the ecophysiological and morphological characteristics of two salt-tolerant tree species, Eucalyptus camaldulensis Dehn. and Combretum quadrangulare Kurz. A greenhouse experiment with different levels of NaCl salinity (0, 0.5, 1.0, 1.5, and 2.0%) was set up and the results were compared with those of a field study on non-saline and saline soils. The determination of optimum gas exchange and the development and evaluation of photosynthetic models with and without water deficit were also included in this study.
Morphological characteristics under saline conditions showed that shoot height and diameter growth, shoot internode length, root length/biomass, leaf width and length, leaf area, number and biomass, and shoot/root and leaf/root ratios decreased with salinity, while leaf thickness increased with salinity. More growth was allocated to the roots than to the leaf canopy. Ecophysiological studies in laboratory showed that photosynthesis, stomatal conductance and water potential decreased with salinity, while the CO2 compensation point increased with salinity. Transpiration, dark respiration and photorespiration increased at low salinity but decreased at high salinity levels. In the field study, however, there were no significant differences in stomatal conductance and opening between saline and non-saline soils. Model predictions supported the results of the field measurements. Adaptation to salinity was reflected in an acclimatization of tree structure in the field study. There were both functioning and structural changes of seedlings in the greenhouse experiment
In terms of ecophysiological and morphological characteristics, E. Camaldulensis showed better salt tolerance than C. Quadragulare both in the greenhouse experiment and field study
The PDF includes a summary in Finnish.
At the beginning of the investigation period the total biomass of the Scots pine (Pinus sylvestris L.) stands on the ordinary sedge pine mire was 48 t/ha. The biomass of the mixed stands of Scots pine and birch (Betula pubescens Erhr.) on the herbrich sedge pine mire was 91 t/ha, out of which 60% was from pine. The biomass of the Norway spruce (Picea abies (L.) H. Karst.) on the Vaccinium-Myrtillus spruce mire was 148 t/ha. The average annual net increment of the stand biomass was 5.8 t/ha in the unfertilized pine stand and 6.7 t/ha in the NPK and micronutrient fertilized one during the six-year investigation period. The corresponding figures in the mixed stand were 7.2 t/ha and 7.6 t/ha. The net increment of the biomass in the unfertilized spruce stand was 6.9 t/ha and in the fertilized 8.4 t/ha. A considerable proportion of the net increment was lost to the ground as litter in all stands.
The nitrogen, phosphorus, potassium, magnesium, iron, manganese, zinc, copper and boron cycles were investigated. The annual nitrogen uptake from the soil was 26–42 kg/ha, that of phosphorus 2.5–3.4 kg/ha, potassium 4.5–12 kg/ha, calcium 12–29 kg/ha, magnesium 2–4 kg/ha, iron 1.4–6.6 kg/ha, manganese less than 2 kg/ha and the other nutrients only some grams. Only part of the fertilized nutrients was fixed in the stand.
The PDF includes a summary in Finnish.