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Articles containing the keyword 'adaptation'.

Category: Commentary

article id 528, category Commentary
Annikki Mäkelä, Thomas J. Givnish, Frank Berninger, Thomas N. Buckley, Graham D. Farquhar, Pertti Hari. (2002). Challenges and opportunities of the optimality approach in plant ecology. Silva Fennica vol. 36 no. 3 article id 528. https://doi.org/10.14214/sf.528
A meeting was held in Hyytiälä, Finland 10–12 April 2000 to assess critically the current challenges and limitations of the optimality approach in plant ecophysiology and botany. This article summarises the general discussions and views of the participants on the use of optimisation models as tools in plant ecophysiological research. A general framework of the evolutionary optimisation problem is sketched with a review of applications, typically involved with balanced regulation between parallel processes. The usefulness and limitations of the approach are discussed in terms of published examples, with special reference to model testing. We conclude that, regardless of inevitable problems of model formulation, wider application of the optimality approach could provide a step forward in plant ecophysiology. A major role of evolutionary theory in this process is simply the formulation of testable hypotheses, the evaluation of which can lead to important advances in our ecophysiological understanding and predictive ability.
  • Mäkelä, University of Helsinki, Dept. of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: annikki.makela@helsinki.fi (email)
  • Givnish, University of Wisconsin, Department of Botany, Madison, WI 53706 USA ORCID ID:E-mail:
  • Berninger, University of Helsinki, Dept. of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Buckley, Cooperative Research Centre for Greenhouse Accounting and Environmental Biology Group, and Research School of Biological Sciences, Australian National University, ACT 2601, Australia ORCID ID:E-mail:
  • Farquhar, Cooperative Research Centre for Greenhouse Accounting and Environmental Biology Group, and Research School of Biological Sciences, Australian National University, ACT 2601, Australia ORCID ID:E-mail:
  • Hari, University of Helsinki, Dept. of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:

Category: Research article

article id 10076, category Research article
Tore Skrøppa, Arne Steffenrem. (2019). Genetic variation in phenology and growth among and within Norway spruce populations from two altitudinal transects in Mid-Norway. Silva Fennica vol. 53 no. 1 article id 10076. https://doi.org/10.14214/sf.10076
Highlights: Norway spruce populations distributed along each of two altitudinal transects showed strong clinal relationships between the annual mean temperatures of the sites of the populations and height and phenology traits in short term tests and height in field trials; Large variation was present among families within populations for height and phenology traits and with a wider range within than among populations; Correlation patterns among traits were different for provenances and families.

Progenies from open pollinated cones collected in natural populations of Norway spruce (Picea abies (L.) Karst.) distributed along two altitudinal transects in Mid-Norway were tested in the nursery, in short term tests and in long-term field trials. The populations showed clinal variation related to the mean annual temperatures of the populations, with the earliest bud flush and cessation of shoot elongation and lowest height at age nine years for the high altitude populations. Within population variation was considerable as the narrow sense heritability for these traits was 0.67, 0.31 and 0.09 in one transect and 0.55, 0.18 and 0.14 in the other transect, respectively. Lammas shoots occurred in the short term trials with large variation in frequency between years. There was significant family variation for this trait, but also interactions between populations and year. The variance within populations was considerably larger in the populations from low altitude compared to the high-altitude populations. Significant genetic correlations between height and phenology traits and damage scores indicate that families flushing early and ceasing growth late were taller. Taller families also had higher frequencies of damages. Selection of the top 20% families for height growth in short term tests at age nine years gave a simulated gain of 11% increased height growth at age 18 years in long term trials at altitudes similar to those of origin of the populations. The gain was negative when high altitude populations were selected based on testing in the lowland.

  • Skrøppa, Norwegian Institute of Bioeconomy Research, P.O. Box 115, 1431 Ås, Norway ORCID ID:E-mail: tore.skroppa@nibio.no (email)
  • Steffenrem, ORCID ID:E-mail:
article id 10040, category Research article
Bengt Andersson Gull, Torgny Persson, Aleksey Fedorkov, Tim J. Mullin. (2018). Longitudinal differences in Scots pine shoot elongation. Silva Fennica vol. 52 no. 5 article id 10040. https://doi.org/10.14214/sf.10040
Highlights: More northerly Scots pine origins exhibit earlier onset and cessation of shoot growth; Continental origins show more northern phenological behaviour; Heat accumulation requirements for onset are not fixed and may be lower when accumulating slower; Scots pine may suffer from spring frost due to earlier growth onset in a warming climate; Phenological traits show potential to adapt to new climate conditions by breeding.

Phenology can have a profound effect on growth and climatic adaptability of long-lived, northern tree species such as Scots pine (Pinus sylvestris L.), where the onset of growth in the spring is triggered mainly by accumulated heat, while cessation of growth is related to the joint effect of photoperiod and temperature. In this study, the objectives were: (1) to compare shoot phenology of genetic material from Scandinavia (maritime climate origin) and northern Russia (continental climate origin) sources, under field conditions in both Scandinavia and Russia (maritime and continental growth conditions); and (2) to estimate the heritabilities of phenological parameters. The material used was part of a larger provenance test series involving Scots pine populations and open-pollinated plus-tree families from Russia, Sweden and Finland. Terminal shoot elongation was measured on multiple occasions during the seventh growing season from seed at a trial near Bäcksjön (Sweden) and Syktyvkar (northern Russia). We calculated the regression of relative shoot elongation over accumulated heat sum above +5 °C using an exponential expression. Seedlings of Swedish and Russian provenance had similar heat-sum requirements for growth onset and cessation in both trials. More northern provenances started onset and cessation at a lower temperature sum, but heat accumulation requirements for onset were not fixed. Scots pine may suffer from spring frost due to earlier growth onset in a warming climate. Variation and heritability of phenological traits show potential to adapt Scots pine to new climate conditions by breeding.

  • Andersson Gull, The Swedish Forestry Research Institute (Skogforsk), Box 3, SE-918 21 Sävar, Sweden ORCID ID: https://orcid.org/0000-0003-3556-3172 E-mail: bengt.anderssongull@skogforsk.se
  • Persson, The Swedish Forestry Research Institute (Skogforsk), Box 3, SE-918 21 Sävar, Sweden ORCID ID:E-mail: torgny.persson@skogforsk.se
  • Fedorkov, The Institute of Biology of Komi Scientific Centre of the Ural Branch of the Russian Academy of Sciences (IB Komi SC UB RAS), Kommunisticheskaya St., 28, Syktyvkar, 167982, Russia ORCID ID: https://orcid.org/0000-0001-7800-7534 E-mail: fedorkov@ib.komisc.ru
  • Mullin, The Swedish Forestry Research Institute (Skogforsk), Box 3, SE-918 21 Sävar, Sweden ORCID ID: https://orcid.org/0000-0003-4924-1836 E-mail: tim.mullin@skogforsk.se (email)
article id 1562, category Research article
Mats Erik Berlin, Torgny Persson, Gunnar Jansson, Matti Haapanen, Seppo Ruotsalainen, Lars Bärring, Bengt Andersson Gull. (2016). Scots pine transfer effect models for growth and survival in Sweden and Finland. Silva Fennica vol. 50 no. 3 article id 1562. https://doi.org/10.14214/sf.1562
Highlights: Scots pine transfer effect models for growth and survival, valid in both Sweden and Finland have been developed; The models use high-resolution gridded climate data and can predict performance in future climatic conditions; The models perform well both for unimproved and genetically improved material and can be used to develop deployment recommendations of contemporary forest regeneration material in Sweden and Finland.

In this study, we developed models of transfer effects for growth and survival of Scots pine (Pinus sylvestris L.) in Sweden and Finland using a general linear mixed-model approach. For model development, we used 378 provenance and progeny trials with a total of 276 unimproved genetic entries (provenances and stand seed check-lots) distributed over a wide variety of climatic conditions in both countries. In addition, we used 119 progeny trials with 3921 selected genetic entries (open- and control pollinated plus-tree families) for testing model performance. As explanatory variables, both climatic indices derived from high-resolution gridded climate datasets and geographical variables were used. For transfer, latitude (photoperiod) and, for describing the site, temperature sum were found to be main drivers for both survival and growth. In addition, interaction terms (between transfer in latitude and site altitude for survival, and transfer in latitude and temperature sum for growth) entail changed reaction patterns of the models depending on climatic conditions of the growing site. The new models behave in a way that corresponds well to previous studies and recommendations for both countries. The model performance was tested using selected plus-trees from open and control pollinated progeny tests. Results imply that the models are valid for both countries and perform well also for genetically improved material. These models are the first step in developing common deployment recommendations for genetically improved forest regeneration material in both Sweden and Finland.

  • Berlin, Uppsala Science Park, SE-75183 Uppsala, Sweden ORCID ID:E-mail: mats.berlin@skogforsk.se (email)
  • Persson, Skogforsk, Box 3, SE-91821 Sävar, Sweden ORCID ID:E-mail: torgny.persson@skogforsk.se
  • Jansson, Uppsala Science Park, SE-75183 Uppsala, Sweden ORCID ID:E-mail: gunnar.jansson@skogforsk.se
  • Haapanen, Natural Resources Institute Finland (Luke), Green Technology, Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail: matti.haapanen@luke.fi
  • Ruotsalainen, Natural Resources Institute Finland (Luke), Green Technology, Finlandiantie 18, FI-58450 Punkaharju, Finland ORCID ID:E-mail: seppo.ruotsalainen@luke.fi
  • Bärring, Rossby Centre, Swedish Meteorological and Hydrological Institute, Folkborgsvägen 17, SE-60176 Norrköping, Sweden ORCID ID:E-mail: lars.barring@smhi.se
  • Andersson Gull, Skogforsk, Box 3, SE-91821 Sävar, Sweden ORCID ID:E-mail: bengt.andersson@skogforsk.se
article id 1274, category Research article
Juho Hautsalo, Paul Mathieu, Sakina Elshibli, Pekka Vakkari, Juha Raisio, Pertti Pulkkinen. (2015). Variation in height and survival among northern populations of pedunculate oak (Quercus robur L.): results of a 13-year field study. Silva Fennica vol. 49 no. 2 article id 1274. https://doi.org/10.14214/sf.1274
Highlights: A height-sum function allowed us to compare the combined growth and survival of northernmost stands of pedunculate oak; Individuals from Turku-Katariinanlaakso performed the best, although other families performed more consistently across trials, which should be considered in future conservation and breeding. Surprisingly, trees planted in a trial location beyond the natural northern limit of pedunculate oak showed the best performance.
We analysed the adaptive potential of pedunculate oak (Quercus robur L.) in terms of variation in height and survival in five field trials located in southern and central Finland. The trials were established with Finnish native material from six different seed origins. Thirteen years after planting, the number of living trees was counted and measured for height. Analysis of height and survival revealed a significant effect of origin, i.e., a genetic basis to individual tree performance. Two origins from the Turku region (Ruissalo and Katariinanlaakso) performed the best while trees originating from Parainen (Lenholmen) performed the worst. In order to study the effects due to tree origin, a comparison of families (half-sibling trees, i.e. those sharing the same ‘mother’ tree) was made by combining height and survival through a height-sum equation (i.e., the product of mean survival and height of each family in each trial) and used to calculate family- and origin-level ecovalences. Ecovalence is a metric for performance consistency, and indicates how much each variable contributes to the total variation; the higher the value, the lower the consistency of trees across the trials based on their origin or family. Analysis of consistency showed similar results to growth and survival, with Turku families performing the best and families from Parainen performing the worst. Families in the Katariinanlaakso stand (Turku) generally had more stable ecovalence values and more dispersed height-sums, while Ruissalo (Turku) families had higher mean height-sum but higher variability in ecovalence values. These results suggest that seed origins (i.e., genotypes) can be optimized in terms of their suitability for commercial or ecological forest management.
  • Hautsalo,  Natural Resources Institute Finland (Luke), Green technology, Antinniementie 1, FI-41330 Vihtavuori, Finland ORCID ID:E-mail: juho.hautsalo@luke.fi (email)
  • Mathieu, Agrocampus Ouest, 35000 Rennes, France ORCID ID:E-mail:
  • Elshibli, University of Helsinki, Helsinki, Finland ORCID ID:E-mail:
  • Vakkari, Natural Resources Institute Finland (Luke), Vantaa, Finland ORCID ID:E-mail: pekka.vakkari@luke.fi
  • Raisio, City of Helsinki, Helsinki, Finland ORCID ID:E-mail:
  • Pulkkinen, Natural Resources Institute Finland (Luke), Vantaa, Finland ORCID ID:E-mail: pertti.pulkkinen@luke.fi
article id 1176, category Research article
Batoul Al-Hawija, Viktoria Wagner, Monika Partzsch, Isabell Hensen. (2014). Germination differences between natural and afforested populations of Pinus brutia and Cupressus sempervirens. Silva Fennica vol. 48 no. 4 article id 1176. https://doi.org/10.14214/sf.1176
Highlights: Silvicultural practices of raising and outplanting seedlings yielded contrasting outcomes in our species; Afforested Pinus brutia populations acquired ability to tolerate drought stress at intermediate and hot temperatures compared to natural populations, which may indicate local adaptation; Natural Cupressus sempervirens populations showed higher salt-tolerance than afforested populations; Seed germination was optimal under intermediate temperatures and deionized water for both species.
In afforestation, silvicultural processes of raising and planting seedlings under certain conditions can yield contrasting outcomes in tree stock performance. Moderate nursery conditions may select against stress tolerance whereas planting seedlings in stressful environments at afforestation sites may select for higher stress tolerance compared to natural populations. We compared germination performance between natural and afforested populations of Pinus brutia Ten. subsp. brutia and Cupressus sempervirens L. var. horizontalis (Mill.) under differing stress treatments. Seeds were collected from both natural stands and from afforested populations outside the natural distribution range, in Syria. Cold, intermediate and hot temperature regimes were simulated (8/4 °C, 20/10 °C and 32/20 °C) along with cold stratification, drought stress (–0.2 and –0.4 MPa), salt stress (50 and 100 mMol l–1), and deionized water (control) conditions. In addition, we tested the effects of seed weight and climatic conditions on seed germination. In general, intermediate temperatures were optimal for both population types. Afforested P. brutia populations outperformed natural ones under drought stress levels at hot and/or intermediate temperatures. Conversely, in C. sempervirens, cold stratification at all temperatures and higher salt stress at intermediate temperatures significantly decreased germination in afforested populations. Seed weight did not significantly affect germination percentages, which were however significantly negatively related to annual precipitation in P. brutia, and to annual temperature in C. sempervirens. We infer that silvicultural processes led to divergent outcomes in our species: local adaptation to drought stress and hot temperatures in afforested P. brutia populations and lower salt-stress tolerance in C. sempervirens.
  • Al-Hawija, Martin-Luther-University Halle-Wittenberg, Institute of Biology/Geobotany and Botanical Garden, Am Kirchtor 1, D-06108 Halle/Saale, Germany ORCID ID:E-mail: batoulh@gmail.com (email)
  • Wagner, Department of Botany and Zoology, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic ORCID ID:E-mail: wagner@sci.muni.cz
  • Partzsch, Martin-Luther-University Halle-Wittenberg, Institute of Biology/Geobotany and Botanical Garden, Am Kirchtor 1, D-06108 Halle/Saale, Germany ORCID ID:E-mail: monika.partzsch@botanik.uni-halle.de
  • Hensen, Martin-Luther-University Halle-Wittenberg, Institute of Biology/Geobotany and Botanical Garden, Am Kirchtor 1, D-06108 Halle/Saale, Germany & German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany ORCID ID:E-mail: isabell.hensen@botanik.uni-halle.de
article id 52, category Research article
Mirja Rantala, Teppo Hujala, Mikko Kurttila. (2012). Measuring and monitoring socio-cultural sustainability in the action of forest biodiversity cooperation networks. Silva Fennica vol. 46 no. 3 article id 52. https://doi.org/10.14214/sf.52
To safeguard overall sustainability in forest resource management, the ecological, economic, social, and cultural dimensions of sustainability should all be considered. However, the socio-cultural impacts are frequently contemplated only weakly in sustainability assessments. Hitherto, attempts to operationalize socio-cultural impacts arising from economic utilization or conservation of forest resources have been perceived as vague when compared to rigorous ecological and economic indicators. One reason is that socio-cultural impacts of forest management on individuals and communities are many and by nature context- and case-specific: they need local definition, which hampers diffusion of good solutions. This study developed a multi-criteria method for measuring and monitoring socio-cultural impacts of forest resource management; the case of cooperation network projects within Forest Biodiversity Programme for Southern Finland (METSO) provided empirical data. Based on a literature review, a set of 10 criteria and 25 indicators was compiled. Cumulative utility scores, presenting networks’ contributions to socio-cultural sustainability, were generated using performance, expert evaluation and weighting data and an additive utility model. The method enables longitudinal monitoring of socio-cultural impacts, which is beneficial because outcomes are different at different time points of projects’ life cycles and some appear with a delay. The method can be used in comparing sub-utility distributions i.e. monitoring units’ performance profiles, providing valuable information for policy-makers. The multi-criteria approach and the list of socio-cultural criteria are internationally transferable to other countries and contexts such as forest bioenergy, nature tourism, watershed management, that call for analysing socio-cultural impacts of forest resource management activity on private lands.
  • Rantala, Finnish Forest Research Institute, Joensuu Unit, P.O. Box 68, FI-80101 Joensuu, Finland ORCID ID:E-mail:
  • Hujala, Finnish Forest Research Institute, Vantaa Unit, Vantaa, Finland ORCID ID:E-mail:
  • Kurttila, Finnish Forest Research Institute, Joensuu Unit, P.O. Box 68, FI-80101 Joensuu, Finland ORCID ID:E-mail: mikko.kurttila@metla.fi (email)
article id 93, category Research article
Pertti Pulkkinen, Saila Varis, Raimo Jaatinen, Aulis Leppänen, Anne Pakkanen. (2011). Increasing survival and growth of Scots pine seedlings with selection based on autumn coloration. Silva Fennica vol. 45 no. 4 article id 93. https://doi.org/10.14214/sf.93
This study evaluates the possibility of using autumn coloration of young Scots pine (Pinus sylvestris L.) seedlings as an indicator of adaptation to harsh climate conditions. One-year old seedlings from natural stands with different origins and seed orchards were classified as “red/reddish” and “green” based on the needle color after artificially increased night length in nursery and then measured after 14 years in field trials. In almost all the studied groups seedlings classified as “red/reddish” had significantly higher survival rate than seedlings classified as “green”. The survival of “red/reddish” was 14.2% higher than “green” among natural stand seed material and 56.2% among seed orchard material. During the study period the survival difference between “red/reddish” and “green” seedlings tended to increase. The seedling color had limited connection with the height growth, even though the trees classified as “red/reddish” were slightly taller than those classified as “green”. However, the total productivity over all field trials, described here as a heightsum, of “red/reddish” trees was 15% higher than productivity of “green” trees from natural stand material, and 61% higher than those from seed orchard material. It seems that controlled selection based on autumn color can be utilized within seed crops of different types with the aim to increase the adaptability of seed material to different environmental conditions.
  • Pulkkinen, Metla, Haapastensyrjä, Läyliäinen, Finland ORCID ID:E-mail: pertti.pulkkinen@metla.fi (email)
  • Varis, Metla, Haapastensyrjä, Läyliäinen, Finland ORCID ID:E-mail:
  • Jaatinen, Metla, Haapastensyrjä, Läyliäinen, Finland ORCID ID:E-mail:
  • Leppänen, Metla, Haapastensyrjä, Läyliäinen, Finland ORCID ID:E-mail:
  • Pakkanen, Metla, Haapastensyrjä, Läyliäinen, Finland ORCID ID:E-mail:
article id 168, category Research article
Saila Varis, Anne Pakkanen, Aina Galofré, Pertti Pulkkinen. (2009). The extent of south-north pollen transfer in Finnish Scots pine. Silva Fennica vol. 43 no. 5 article id 168. https://doi.org/10.14214/sf.168
In order to evaluate the possibility of long distance gene flow in Scots pine (Pinus sylvestris L.), we measured the amount and germinability of airborne pollen and flowering phenology in central, northern, and northernmost Finland during 1997–2000. Totally 2.3% of the detected germinable pollen grains were in the air prior to local pollen shedding. The mean number of germinable pollen grains m–3 air per day was lower prior to local pollen shedding, but in the year 2000 there were more germinable pollen grains in the air of central study site prior to local pollen shedding. Prior to the onset of pollen shedding, 7.5% of female strobili which we observed were receptive. On average female strobili became receptive three days earlier than local pollen shedding started. During the period of pollen shedding in the central study site, we detected germinable airborne pollen in the northern site in years 1997, 1999 and 2000. At the northermost site, we detected germinable airborne pollen during the pollen-shedding period of the northern site in 2000. Our detection of germinable airborne pollen and synchrony of strobili maturation from south to north suggest that populations of Scots pine in central and northern Finland may provide genetic material to populations in northern and northernmost Finland, respectively.
  • Varis, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail: saila.varis@metla.fi (email)
  • Pakkanen, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail:
  • Galofré, Passeig de l’estació 21, 5-1, 43800 Valls, Tarragona, Spain ORCID ID:E-mail:
  • Pulkkinen, Finnish Forest Research Institute, Haapastensyrjä Breeding Station, Karkkilantie 247, FI-12600 Läyliäinen, Finland ORCID ID:E-mail:
article id 226, category Research article
Anneli Viherä-Aarnio, Pirkko Velling. (2008). Seed transfers of silver birch (Betula pendula) from the Baltic to Finland – effect on growth and stem quality. Silva Fennica vol. 42 no. 5 article id 226. https://doi.org/10.14214/sf.226
Silver birch (Betula pendula Roth) seed origins from the Baltic countries, Finland and Russia were compared for survival, growth and stem quality, and the effect of latitudinal seed transfer distance examined in two provenance trials. The trials were located on moist upland forest soils at Tuusula (60°21’N) in southern Finland and at Viitasaari (63°11’N) in central Finland. The material consisted of 21 stand and single tree origins ranging from latitudes 54° to 63°N. Survival, height, dbh, relative stem taper, stem volume/ha and the proportion of trees with a stem defect (vertical branch or forked stem), were assessed when the trees were 22 years old. Significant differences were detected among the origins regarding all the measured traits in both trials. Southern Finnish origins produced the highest volume per unit area in central Finland, whereas Estonian and north Latvian stand seed origins, as well as the southern Finnish plus tree origins, were the most productive ones in southern Finland. The more southern the origin, the higher was the proportion of trees with a stem defect in both trials. The latitudinal seed transfer distance had a significant but relatively small effect on survival, stem volume/ha and proportion of trees with a stem defect. The proportion of trees with a stem defect increased linearly in relation to the seed transfer distance from the south. The relationship of both survival and stem volume/ha to the seed transfer distance was curvilinear. Volume/ha was increased by transferring seed from ca. 2 degrees of latitude from the south. A longer transfer from the south, as well as transfer from the north, decreased the yield.
  • Viherä-Aarnio, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail: anneli.vihera-aarnio@metla.fi (email)
  • Velling, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail:
article id 634, category Research article
Marcus Lindner, Petra Lasch, Markus Erhard. (2000). Alternative forest management strategies under climatic change – prospects for gap model applications in risk analyses. Silva Fennica vol. 34 no. 2 article id 634. https://doi.org/10.14214/sf.634
The projected global climate change will influence growth and productivity of natural and managed forests. Since the characteristics of the future regional climate are still uncertain and the response of our forests to changes in the atmospheric and climatic conditions may be both positive or negative, decision making in managed forests should consider the new risks and uncertainties arising from climatic change, especially if the rotation periods are long. An extended version of the forest gap model FORSKA was applied to simulate the forest development at 488 forest inventory plots in the federal state of Brandenburg, Germany, under two climate and three management scenarios. The transient growth dynamics from 1990 to 2100 were investigated at four sites in different parts of the state, representing the variability of environmental and forest conditions within Brandenburg. The alternative management strategies led to distinct differences in forest composition after 110 years of simulation. The projected climate change affected both forest productivity and species composition. The impacts of alternative management scenarios are discussed. It is concluded that the extended forest gap model can be a valuable tool to support decision making in forest management under global change.
  • Lindner, Potsdam Institute for Climate Impact Research, P.O. Box 60 12 03, D-14412 Potsdam, Germany ORCID ID:E-mail: lindner@pik-potsdam.de (email)
  • Lasch, Potsdam Institute for Climate Impact Research, P.O. Box 60 12 03, D-14412 Potsdam, Germany ORCID ID:E-mail:
  • Erhard, Potsdam Institute for Climate Impact Research, P.O. Box 60 12 03, D-14412 Potsdam, Germany ORCID ID:E-mail:

Category: Review article

article id 313, category Review article
Heikki Hänninen, Koen Kramer. (2007). A framework for modelling the annual cycle of trees in boreal and temperate regions. Silva Fennica vol. 41 no. 1 article id 313. https://doi.org/10.14214/sf.313
Models of the annual development cycle of trees in boreal and temperate regions were reviewed and classified on the basis of their ecophysiological assumptions. In our classification we discern two main categories of tree development: 1) fixed sequence development, which refers to irreversible ontogenetic development leading to visible phenological events such as bud burst or flowering, and 2) fluctuating development, which refers to reversible physiological phenomena such as the dynamics of frost hardiness during winter. As many of the physiological phenomena are partially reversible, we also describe integrated models, which include aspects of both fixed-sequence and fluctuating development. In our classification we further discern simple E-models, where the environmental response stays constant, and more comprehensive ES-models, where the environmental response changes according to the state of development. On the basis of this model classification, we have developed an operational modelling framework, in which we define an explicit state variable and a corresponding rate variable for each attribute of the annual cycle considered. We introduce a unifying notation, which we also use when presenting a selection of previously published models. To illustrate the various developmental phenomena and their modelling, we have carried out model simulations. Finally, we discuss the ecophysiological interpretation of the model variables, methodological aspects of the empirical development and testing of the models, the introduction of new aspects to the modelling, other closely related models, and applications of the models.
  • Hänninen, Plant Ecophysiology and Climate Change Group (PECC), Department of Biological and Environmental Sciences, Box 65, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: heikki.hanninen@helsinki.fi (email)
  • Kramer, Alterra, P.O. Box 47, 6700 AA Wageningen, The Netherlands ORCID ID:E-mail:
article id 489, category Review article
Ralph J. Alig. (2003). U.S. landowner behavior, land use and land cover changes, and climate change mitigation. Silva Fennica vol. 37 no. 4 article id 489. https://doi.org/10.14214/sf.489
Landowner behavior is a major determinant of land use and land cover changes, an important consideration for policy analysts concerned with global change. Study of landowner behavior aids in designing more effective incentives for inducing land use and land cover changes to help mitigate climate change by reducing net greenhouse gas emissions. Afforestation, deforestation, reforestation, and timber harvest are the most frequent land management practices that influence forest carbon stocks and flux. Research studies provide estimates of how private landowners respond to market signals and government programs and how they alter land management. For example, landowners have tended to retain subsidized afforested stands well beyond program life in the United States, suggesting that similar programs for climate change mitigation could result in high rates of retention. At the same time, policy makers need to be aware that unintended consequences of policies can lead to significantly different outcomes than envisioned, including leakage possibilities.
  • Alig, USDA Forest Service, Forestry Sciences Lab, 3200 SW Jefferson Way, Corvallis, Oregon 97331, USA ORCID ID:E-mail: ralig@fs.fed.us (email)

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. https://doi.org/10.14214/sf.534
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 ORCID ID:E-mail: leinonen@ou.edu (email)
  • Hänninen, University of Helsinki, Department of Ecology and Systematics, FIN-00014 Helsinki, Finland ORCID ID:E-mail:
article id 609, category Research note
Aleksei Fedorkov. (2001). Climatic adaptation of seed maturity in Scots pine and Norway spruce populations. Silva Fennica vol. 35 no. 1 article id 609. https://doi.org/10.14214/sf.609
Seed maturation of Scots pine and Norway spruce in a provenance experiment at Kortkeros (northern Russia) was examined by the X-ray method. Logarithmic relationships were found between seed anatomy development and long-term average thermal sum. Seed development in the northern populations of Scots pine and Norway spruce was a little faster than in the southern ones.
  • Fedorkov, Russian Academy of Sciences, Komi Science Centre, Institute of Biology, Kommunisticheskya St., 28, 167610 Syktyvkar, Russia ORCID ID:E-mail: fedorkov@ib.komisc.ru (email)

Category: Article

article id 5582, category Article
Jonathan J. Ruel, Matthew P. Ayres. (1996). Variation in temperature responses among populations of Betula papyrifera. Silva Fennica vol. 30 no. 2–3 article id 5582. https://doi.org/10.14214/sf.a9227

How will global warming affect southern populations of boreal trees? In paper birch, Betula papyrifera (Betulaceae), alpine trees with an evolutionary history of relatively cool summers may be more sensitive to climate warming than valley populations. We evaluated this scenario by growing seedlings from different populations in four temperature treatments (mountain field site, valley field site, and two greenhouse rooms).

Populations from low elevations germinated earlier and had higher germination success than population from high elevations (16.8 vs. 22.0 d; 72% vs. 11%). At the valley site, seedlings from native populations grew faster than seedlings from higher elevations (mean ± SE = 0.25 ± 0.02 vs. 0.09 ± 0.04 mm · cm-1 · d-1) while at the mountain site, all seedlings grew at similar rates. Seedling grown in cooler environments had higher root : shoot ratios, perhaps to compensate for temperature limitations in nutrient uptake by roots. Leaf area varied among populations but was not affected by environmental differences across the field sites. Net photosynthetic rates at valley temperatures were higher for seedlings grown in the valley than for seedling grown in the mountains or the warm greenhouse (12.0 vs. 10.3 and 5.8 μmoles · m-2 · s-1), perhaps due to adaptive phenotypic adjustments. Climatic warming could rapidly produce important phenotypic changes in birch trees (e.g. decreased root : shoot ratio, reduced growth in alpine populations). On a longer time-scale, warming could also result in genetic changes as natural selection favours valley genotypes in alpine sites where they are presently rare.

  • Ruel, ORCID ID:E-mail:
  • Ayres, ORCID ID:E-mail:
article id 5558, category Article
Kari Tuomela, Markku Kanninen. (1995). Effects of vapour pressure deficit and soil water content on leaf water potential between selected provenances of Eucalyptus microtheca in an irrigated plantation, eastern Kenya. Silva Fennica vol. 29 no. 3 article id 5558. https://doi.org/10.14214/sf.a9209

The aim of the study was to compare the behaviour of three selected provenances of Eucalyptus microtheca F. Muell. that were likely to respond differently to drought. For this purpose, we studied the effects of vapour pressure deficit and soil water content on leaf water potential in an irrigated plantation in Bura, eastern Kenya.

An international provenance trial of Eucalyptus microtheca, established as a part of Finnida-supported Bura Forestry Research Project in eastern Kenya in 1984 was used as a plant material in the study. The eastern provenance showed generally the lowest leaf water potential on a daily basis. Statistically significant differences in the daily leaf water potential fluctuations were detected. The eastern provenance exhibited the greatest and the northern one the smallest values. The minimum daily leaf water potential of the provenances responded well to changes in gravimetric soil water content, the western provenance being the most sensitive one. The relationship of the observed results and annual rainfall distribution in the geographic regions of the studied provenances is discussed.

  • Tuomela, ORCID ID:E-mail:
  • Kanninen, ORCID ID:E-mail:
article id 5362, category Article
Seppo Kellomäki, Heikki Hänninen, Taneli Kolström. (1988). Model computations on the impacts of the climatic change on the productivity and silvicultural management of the forest ecosystem. Silva Fennica vol. 22 no. 4 article id 5362. https://doi.org/10.14214/sf.a15519

The model computations indicate that the climatic change in the form of higher temperatures and more precipitation could increase the productivity of the forest ecosystem and lead to higher rates of regeneration and growth. More frequent and intensive thinnings are needed to avoid the mortality of trees induced by accelerated maturation and attacks of fungi and insects. The climatic change could support the dominance of deciduous tree species and necessitate an intensification of the tending of seedling stands of conifers. The rise of air temperature during autumn and winter could change also the annual growth rhythm of trees and result in dehardening and subsequent frost damages and attacks of insects and fungi. The pest management could be the greatest challenge to the future silviculture, which could be modified most in Northern Finland.

The PDF includes an abstract in Finnish.

  • Kellomäki, ORCID ID:E-mail:
  • Hänninen, ORCID ID:E-mail:
  • Kolström, ORCID ID:E-mail:
article id 5257, category Article
Heikki Hänninen. (1986). Metsäpuiden vuosirytmitutkimuksen käsitteistä ja teorioista. Silva Fennica vol. 20 no. 1 article id 5257. https://doi.org/10.14214/sf.a15436
English title: Conceptual remarks about the study of the annual rhythm of forest trees.

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.

The PDF includes a summary in English.

  • Hänninen, ORCID ID:E-mail:
article id 5160, category Article
P. M. A. Tigerstedt. (1982). Metsäpuiden populaatiogenetiikka. Helsingissä 1981 pidetyn symposion tutkimusraportit. Silva Fennica vol. 16 no. 2 article id 5160. https://doi.org/10.14214/sf.a15075
English title: Population genetics of forest trees.

The publication comprises proceedings of a conference held in Helsinki in 1981. Forest tree populations are investigated for population genetic structure, mating systems, mechanisms of genetic adaptation and ecological adaptation. Methods and techniques used in population genetic research of forest trees are presented. Much concern is given to applications by means of forest tree breeding, particularly the seed orchard breeding technique. Generally, the application of population genetics in cultivated forests is discussed.

The PDF includes a preface and the presentations of the conference (25 short papers) in English, and a comprehensive summary of the themes of the conference in Finnish.

  • Tigerstedt, ORCID ID:E-mail:
article id 7523, category Article
Alpo Luomajoki. (1999). Differences in the climatic adaptation of silver birch (Betula pendula) and downy birch (B. pubescens) in Finland based on male flowering phenology. Acta Forestalia Fennica no. 263 article id 7523. https://doi.org/10.14214/aff.7523

Male flowering was studied at the canopy level in 10 silver birch (Betula pendula Roth) stands from 8 localities and 14 downy birch (B. pubescens Ehrh.) stands from 10 localities in Finland in 1963–73. Distribution of cumulative pollen catches was compared to the normal Gaussian distribution. The basis for timing of flowering was the 50% point of the anthesis-fitted normal distribution. To eliminate effects of background pollen, only the central, normally distributed part of the cumulative distribution was used. Development was measured and tested in calendar days, in degree days (> 5°C) and in period units. The count of the parameters began in March 19.

Male flowering in silver birch occurred from late April to late June depending on latitude, and flowering in downy birch took place from early May to early July. The heat sums needed for male flowering varied in downy birch stands latitudinally but there was practically no latitudinal variation in silver birch flowering. The amount of male flowering in stands of the both species were found to have a large annual variation but without any clear periodicity.

The between years pollen catch variation in stands of either birch species did not show any significant latitudinal correlation in contrast to Norway spruce stands. The period unit heat sum gave the most accurate forecast of the timing of flowering for 60% of the silver birch stands and for 78.6% of the downy birch stands. Silver birch seems to have a local inclination for a more fixed flowering date compared to downy birch, which could mean a considerable photoperiodic influence on flowering time of silver birch. The species had different geographical correlations.

Frequent hybridization of the birch species occurs more often in Northern Finland than in more southerly latitudes. The different timing in the flowering causes increasing scatter in flowering times in the north, especially in the case of downy birch. Thus, the change of simultaneous flowering of the species increases northwards due to a more variable climate and higher altitudinal variation. Compared with conifers, the reproduction cycles of the two birch species were found to be well protected from damage by frost.

  • Luomajoki, ORCID ID:E-mail:
article id 7504, category Article
Alpo Luomajoki. (1993). Kuusen sopeutuminen Suomen ilmastoon hedekukkimisaikojen perusteella arvioituna. Acta Forestalia Fennica no. 242 article id 7504. https://doi.org/10.14214/aff.7504
English title: Climatic adaptation of Norway spruce (Picea abies (L.) Karsten) in Finland based on male flowering phenology.

Anthesis was studied at the canopy level in 10 Norway spruce (Picea abies (L.) H. Karst.) stands from 9 localities in Finland was studied in 1963-74. Distribution of pollen catches were compared with the normal Gaussian distribution. The basis for the timing studies was the 50% point of the anthesis-fitted normal distribution. Development was characterized in calendar days, in degree days (>5°C) and in period units. The count of each unit began on March 19 (included). Male flowering in Norway spruce stands was found to have more annual variation in quantity than in Scots pine (Pinus sylvestris L.) stands studied earlier.

Anthesis in spruce in Northern Finland occurred at a later date than in the south. The heat sums needed for anthesis varied latitudinally less in spruce than in pine. The variation of pollen catches in spruce increased towards north-west as in the case of Scots pine. In the unprocessed data, calendar days were found to be the most accurate forecast of anthesis in Norway spruce. Locally, the period unit could be a more accurate parameter for the stand average. However, on a calendar day basis, when annual deviations between expected and measured heat sums were converted to days, period units were narrowly superior to days.

The geographical correlations respected to timing of flowering, calculated against distances measured along simulated post-glacial micgation routes, were stronger than purely latitudinal correlations. Effects of the reinvasion of Norway spruce into Finland are thus still visible in spruce populations just as they were in Scots pine populations.

The proportion of the average annual heat sum needed for spruce anthesis grew rapidly north of a latitude of ca. 63° and the heat sum needed for anthesis decreased only slightly towards the timberline. In light of flowering phenology, it seems probable that north-western third of Finnish Norway spruce populations are incompletely adapted to the prevailing cold climate. A moderate warming of the climate would therefore be beneficial for Norway spruce. This accords roughly with the adaptive situation in Scots pine

The PDF includes a summary in Finnish.

  • Luomajoki, ORCID ID:E-mail:
article id 7683, category Article
Alpo Luomajoki. (1993). Climatic adaptation of Scots pine (Pinus sylvestris L.) in Finland based on male flowering phenology. Acta Forestalia Fennica no. 237 article id 7683. https://doi.org/10.14214/aff.7683

Timing of anthesis in 21 Scots pine (Pinus sylvestris L.) stands from 14 localities in Finland was studied at the canopy level in 1963-74. Distribution of pollen catches were compared with the normal Gaussian distribution. The basis for the timing studies was the 50 per cent point of the anthesis-fitted normal distribution. Development was characterized in calendar days, in degree days (>5°C) and in period units. The count of each unit began on March 19 (included).

Period unit was found to be the most accurate delineation of development. Locally, calendar days were sometimes a more accurate parameter. Anthesis in Northern Finland occurred at a later date than in the south as was expected, but at lower heat sum. The variation in the timing of anthesis and the variation of pollen catches increased northwards. The geographical correlations calculated against distances measured along simulated post-glacial migration routes were stronger than purely latitudinal correlations. Effects of the reinvasion of Scots pine into Finland are thus still visible in pine populations.

The proportion of the average annual heat sum needed for anthesis grew rapidly above a latitude of 63° even though the heat sum needed for anthesis decreased towards the timberline. In light of flowering phenology, it seemed probable that the northern populations in Scots pine in Finland have still not completely adapted to the prevailing cold climate at these latitudes. A moderate warming of the climate would therefore be beneficial for Scots pine.

The PDF includes a summary in Finnish.

  • Luomajoki, ORCID ID:E-mail:
article id 7660, category Article
Heikki Hänninen. (1990). Modelling bud dormancy release in trees from cool and temperate regions. Acta Forestalia Fennica no. 213 article id 7660. https://doi.org/10.14214/aff.7660

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.

The PDF includes a summary in Finnish.

  • Hänninen, ORCID ID:E-mail:

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