The relationships between bud dormancy and frost hardiness were examined using two-year-old Pinus sylvestris L. seedlings. The chilling temperatures used were +4 and -2°C. To examine the dormancy release of the seedlings, a forcing technique was used. Frost hardiness was determined by artificial freezing treatments and measurements of electrical impedance. At the start of the experiment, the frost hardiness of the seedlings was about -25°C. After the rest break, the seedlings kept at +4°C dehardened until after eight weeks their frost hardiness reached -5°C. At the lower chilling temperature (-2°C) the frost hardiness remained at the original level. When moved from +4 to -2°C, seedlings were able to reharden only after the time required for bud burst in the forcing conditions had reached the minimum.
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Models concerning the effects of temperature on dormancy release in woody plants were tested using two-year old seedlings of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.). Chilling experiments suggest that the rest period has a distinct end point. Before the attainment of this end point high temperatures do not promote bud development towards dormancy release, and after it further chilling does not affect the subsequent bud development. A new hypothesis of dormancy release is suggested on the basis of a comparison between present and earlier findings. No difference in the proportion of growth commencing seedlings were detected between the forcing temperatures of 17°C and 22°C. The rest break of 50% of Norway spruce and Scots pine seedlings required six and eight weeks of chilling, respectively. Great variation in the chilling requirement was found, especially for Scots pine.
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This issue of Silva Fennica consists of eight articles, which are based on a co-nordic conference ”Frost hardiness and over-wintering in forest tree seedlings”, held in Joensuu, Finland, during December 1–3, 1986. The whole annual cycle of the trees is considered. Emphasis is given on methods for the study of frost hardiness, genetic variation in frost hardiness, nitrogen metabolism, bud dormancy release, and joint effect of natural and anthropogenic stress factors in the winter damage of forest trees. Practical implications for tree breeding and nursery management are discussed.
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The distribution of the minimum temperatures after the beginning of the active period (one temperature for each spring) have been calculated for each principle using daily meteorological data collected during the years 1883–1980. The efficiency criterion is the variance of the minimum temperature distributions and the length of the active period. The most efficient regulation principle is found to be based on the temperature sum which includes a feedback component.
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Logical structure of three simulation models and one conceptual model concerning effects of temperature on dormancy release in woody plants was examined. The three basic types of simulation models differed in their underlying assumptions. Contrasting implications of the models were inferred by deduction. With the aid of these implications, the model types can be tested using experiments with continuous and interrupted chilling. Similarly, implications of the conceptual model of rest phases were inferred, by which the model can be tested using experiments with continuous chilling and forcing in multiple temperatures. The possibilities to synthetize the conceptual model with any of the three simulation model types, as well as the biological interpretation of the model variables, were discussed.
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Different approaches to the study of the annual rhythm of forest trees are described and compared by analysing the concepts and theories presented in the literature. The seasonality varying morphological and physiological state of forest trees is referred to as the annual rhythm s. lat., from which the annual ontogenetic rhythm is separated as a distinct type. The dormancy phenomena of the trees are grouped into four categories. Theories concerning the regulation of the annual rhythm are divided into two main types, the most common examples of which are the photoperiod theory and the temperature sum theory. Recent efforts towards a synthetic theory are described.
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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 premises of several models obtained from literature on bud dormancy release in trees from cool and temperate regions differs from each other with respect to responses to air temperature during the rest period of the buds. The predicted timing of bud burst in natural conditions varied among the models, as did the prediction of the models for the outcome of a chilling experiment.
Experimental results with two-year old seedlings of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.) did not agree with any of the models. The experimental results also deviated from abundand earlier findings, which also disagreed with any of the models. This finding suggests that Finnish provenances of Scots pine and Norway spruce differ from more southern provenances with respect to temperature regulation of bud dormancy release.
A synthesis model for the effects of air temperature on bud dormancy release in trees was developed on the basis of the previous models and the experimental results of both the present and previous studies. The synthesis model contains part of the original models as special cases. The parameters of the synthesis model represent several aspects of the bud dormancy release of trees that should be addressed separately with each species and provenance in experimental studies. Further aspects of dormancy release were discussed, in order to facilitate further development of the models.
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The dependence of the rates of photosynthesis and transpiration are studied on the environmental factors and on the control of the plant metabolism with Scots pine (Pinus sylvestris L.) in the natural environment. The importance of the different environmental factors to the photosynthetic production are evaluated. In modelling the dependence of the rates of photosynthesis and transpiration on the environmental factors and on the control processes, a dynamic system analysis approach is applied. Irradiance, temperature and water are explored during the annual cycle. Field measurements of the CO2 exchange and environmental factors over three years are used in the analysis of the rate of the photosynthesis. The effect of different environmental factors on photosynthetic production is evaluated by model estimations using weather data in a 20-year period.
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The timing of the tetrad phase of microsporogenesis in sixteen tree species, belonging to the genera of Abies, Larix, Picea, Pinus, Alnus, Betula, Corylus and Populus, was studied. The tetrad phase of microsporogenesis in conifers and in Populus tremula L. was reached from late March to early June including the yearly and latitudinal variation. The tetrad phase in Betulaceae was reached in late July to mid-August. The microsporogenesis in Betulaceae species differed in ecophysiological terms from the other species studied in that the timing in Betulaceae was rather day-length dependent than heat sum-correlated. In conifers and in Populus the timing of tetrad phase correlated with heat sums accumulated and did not correlate with day length or any kind of thermal threshold. This difference was, however, judged to be associated to seasonal adaptive strategies rather than taxonomic relationships.
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In this paper a precise definition is given of the term physiological clock and the role of this clock in biological developmental and growth processes is mathematically studied. The heat sum method employed in the study of the annual cycle of development of forest trees has been used as the starting point. The mathematical principles of this method are analysed and it is shown that, on the same principles, a fairly general physiological clock can be constructed. Also, two growth models are presented in which this generalized physiological clock proves to play an important role.
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