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Articles containing the keyword 'phenology'.

Category: Research article

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 5656, category Research article
Lars-Göran Stener, Johan Westin. (2017). Early growth and phenology of hybrid aspen and poplar in clonal field tests in Scandinavia. Silva Fennica vol. 51 no. 3 article id 5656. https://doi.org/10.14214/sf.5656
Highlights: Growth and survival was in general higher for hybrid aspen than for poplar; The poor performance of poplar was likely due to poor climatic adaptation or to high soil acidity; The genetic results indicate good possibilities for effective clonal selection; The results support current recommendations for utilization of selected hybrid aspen and poplar regeneration material in Sweden.

Results on early survival, growth and shoot phenology of hybrid aspen (Populus tremula L. × P. tremuloides Michx.) and poplar clones (P. trichocarpa Torr. & A. Gray, P. balsamifera L., P. maximowiczii A. Henry and their hybrids) in 13 Scandinavian field trials are presented. The trials were established on forest land (7 sites) or former agricultural land (6 sites) within the latitude range of 56° to 65° N and were assessed 3–4 years after establishment. The main aim was to evaluate phenotypic and genetic differences related to early survival, growth and phenology for hybrid aspen and poplar for different site types and latitudes. Growth and survival was generally higher for hybrid aspen than poplar at all sites. The poor performance of poplar compared to hybrid aspen is likely due to climatic maladaptation or high soil acidity. The early growth performance of the species need to be confirmed at a higher age. The genetic variation and genetic control for growth, phenology and survival was in general intermediate to large indicating good possibilities for effective clonal selection. The genetic site x site correlations (rGE) for growth were for hybrid aspen mostly strong, indicating a weak genotype by environment interaction, while rGE were inconsistent for poplars.The result suggests that southern Sweden can be treated as a single test and utilization zone and in northern Sweden the region along the coast may be another zone. It is too early to make any corresponding conclusions for poplar. In addition, the result backs up the current recommendations for utilization of selected hybrid aspen and poplar regeneration material in Sweden.

  • Stener, The Forestry Research Institute of Sweden, Ekebo 2250, 268 90 Svalöv, Sweden ORCID ID:E-mail: lars-goran.stener@skogforsk.se (email)
  • Westin, The Forestry Research Institute of Sweden, Box 3, 918 21 Sävar, Sweden ORCID ID:E-mail: johan.westin@skogforsk.se
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 600, category Research article
Qibin Yu, P. M. A. Tigerstedt, Matti Haapanen. (2001). Growth and phenology of hybrid aspen clones (Populus tremula L. x Populus tremuloides Michx.). Silva Fennica vol. 35 no. 1 article id 600. https://doi.org/10.14214/sf.600
Height, basal diameter, diameter at breast height, bud burst, and leaf development were recorded in a 5-year-old hybrid aspen clonal trial. The field trial consisted of four aspen hybrid clones (Populus tremula x Populus tremuloides) and one local P. tremula seedling source. Phenological traits were observed in the 3rd year. Growth patterns were recorded during the 3rd and 4th years. Phenological traits were explored in relation to hybrid vigor expressed as growth traits. Differences were observed for phenological and growth traits among hybrid clones and P. tremula. The growth period varied from 143–158 days for the four hybrid clones, and was 112 days for P. tremula. The correlation between growth period and yield was highly significant. The annual growth rate of height for the hybrids was 4.2 cm per 7 days (2.4 for P. tremula) in the 3rd year and 6.4 cm per 7 days (2.9 for P. tremula) in the 4th year. After 5 years, mean estimated stem volume of the hybrids was 3.9 times that of P. tremula. Significant clone by year interaction was observed for height, diameter, and volume growth. The hybrid vigor seems to be mainly attributable to a longer growth period.
  • Yu, Department of Plant Biology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: qibin.yu@helsinki.fi (email)
  • Tigerstedt, Department of Plant Biology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Haapanen, Finnish Forest Research Institute, P.O. Box 18, FIN-01301 Vantaa, Finland 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 535, category Review article
Thomas J. Givnish. (2002). Adaptive significance of evergreen vs. deciduous leaves: solving the triple paradox. Silva Fennica vol. 36 no. 3 article id 535. https://doi.org/10.14214/sf.535
Patterns in the dominance of evergreen vs. deciduous plants have long interested ecologists, biogeographers, and global modellers. But previous models to account for these patterns have significant weaknesses. Bottom-up, mechanistic models – based on physiology, competition, and natural selection – have often been non-quantitative or restricted to a small range of habitats, and almost all have ignored belowground costs and whole-plant integration. Top-down, ecosystem-based models have succeeded in quantitatively reproducing several patterns, but rely partly on empirically derived constants and thresholds that lack a mechanistic explanation. It is generally recognized that seasonal drought can favor deciduous leaves, and that infertile soils can favor long-lived evergreen leaves. But no model has yet explained three great paradoxes, involving dominance by 1) evergreens in highly seasonal, boreal forests, 2) deciduous larch in many nutrient-poor peatlands, and 3) evergreen leaf-exchangers in nutrient-poor subtropical forests, even though they shed their leaves just as frequently as deciduous species. This paper outlines a generalized optimality model to account for these and other patterns in leaf longevity and phenology, based on maximizing whole-plant carbon gain or height growth, and building on recent advances in our understanding of the quantitative relationships of leaf photosynthesis, nitrogen content, and mass per unit area to leaf life-span. Only a whole-plant approach can explain evergreen dominance under realistic ecological conditions, or account for the boreal paradox, the larch paradox, the leaf-exchanger paradox, and expected shifts in shade tolerance associated with leaf phenology. Poor soils favor evergreens not merely by increasing the costs of nutrient acquisition, but also by depressing the maximum rate of photosynthesis and thus the seasonal contrast in photosynthetic return between leaves adapted to favorable vs. unfavorable conditions. The dominance of evergreens in western North America beyond the coastal zone of mild winters and winter rainfall appears related to the unusually long photosynthetic season for evergreen vs. deciduous plants there. Future models for optimal leaf phenology must incorporate differences between evergreen and deciduous plants in allocation to photosynthetic vs. non-photosynthetic tissue, rooting depth, stem allometry, xylem anatomy, and exposure to herbivores and leaching, and analyze how these differences interact with the photosynthetic rate, transpiration, and nutrient demands of leaves with different life-spans to affect rates of height growth in specific microsites.
  • Givnish, Dept of Botany, University of Wisconsin, Madison, WI 53706, USA ORCID ID:E-mail: givnish@facstaff.wisc.edu (email)
article id 693, category Review article
Erik G. Ståhl. (1998). Changes in wood and stem properties of Pinus sylvestris caused by provenance transfer. Silva Fennica vol. 32 no. 2 article id 693. https://doi.org/10.14214/sf.693
Wood properties focused in forest tree breeding should be of economic importance, have a large total variation and a high heritability. The properties of interest are those that influence the strength and durability of sawn products or the amount and properties of pulp produced. The following wood properties are treated: width of the annual ring, juvenile wood, late wood content, heart wood, tracheid dimensions, basic density, stem straightness and branch diameter. The provenance variation in wood properties can be related to differences in growth phenology. In the northern part of distribution P. sylvestris (L.) provenances transferred a few degrees southwards have a high survival and yield but stem wood production is low. Trees from these provenances will be straight and with few spike knots or other injuries. The shoot elongation period will be short and the temperature sum required for wood formation sufficient. Provenances transferred southwards will form thin annual rings, few and thin branches, little early wood, high basic density and slender tracheids with thick cell walls in comparison to local provenances. An example of the effect of alternative transfers on the yield and wood properties is evaluated. In regions with deviating climatic patterns alternative provenance transfer patterns may be better. The objectives of the land owner should influence the provenance choice. The importance of integrating tree improvement with silvicultural management is discussed with reference to spacing.
  • Ståhl, College of Dalarna, CITU Centre for Industrial Technology and Development, S-781 88 Borlänge, Sweden ORCID ID:E-mail: esl@du.se (email)

Category: Research note

article id 1443, category Research note
Jouni Partanen, Risto Häkkinen, Heikki Hänninen. (2016). Significance of the root connection on the dormancy release and vegetative bud burst of Norway spruce (Picea abies) seedlings in relation to accumulated chilling. Silva Fennica vol. 50 no. 1 article id 1443. https://doi.org/10.14214/sf.1443
Highlights: Cutting the root connection slightly increased the number of days to bud burst of Norway spruce seedlings under warm conditions but it had no consistent effect on bud burst percentage; Our results obtained with seedlings suggest that using detached tree material in dormancy release experiments may slightly affect the results but it will evidently not lead to drastically erroneous conclusions.

The effect of cutting the root connection by detaching the shoot from the root system on dormancy release and vegetative bud burst was examined in 2-year-old seedlings of Norway spruce (Picea abies [L.] Karst.). Seedlings were transferred at 1–4 week intervals between October and January from outdoor conditions to experimental forcing in a heated greenhouse. Before forcing, half of the seedlings were cut above ground line, and the detached shoots were forced with their cut ends placed in water. The intact seedlings were forced with their root system remaining intact in the pots. Vegetative bud burst was observed visually. Cutting the root connection slightly increased days to bud burst in the forcing conditions, however, no consistent effect on bud burst percentage was found. Our preliminary seedling data suggest that using detached tree material in dormancy release experiments may have a small effect on bud burst date but it will evidently not lead to drastically erroneous conclusions. Further studies, using different seed lots, are needed to assess the effect of detaching on the dormancy release and bud burst, especially in adult trees.

  • Partanen, Natural Resources Institute Finland (Luke), Management and Production of Renewable Resources, Juntintie 154, FI-77600 Suonenjoki, Finland ORCID ID:E-mail: jouni.partanen@luke.fi (email)
  • Häkkinen, Natural Resources Institute Finland (Luke), Management and Production of Renewable Resources, P.O. Box 18, FI-01301 Vantaa, Finland ORCID ID:E-mail: risto.hakkinen@luke.fi
  • Hänninen, University of Helsinki, Department of Biosciences, Viikki Plant Science Centre (ViPS), P.O. Box 65, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: heikki.hanninen@helsinki.fi
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 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. https://doi.org/10.14214/sf.608
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 ORCID ID:E-mail: jouni.partanen@metla.fi (email)
  • Leinonen, University of Joensuu, Faculty of Forestry, P.O. Box 111, FIN-80101 Joensuu, Finland ORCID ID:E-mail:
  • Repo, University of Joensuu, Faculty of Forestry, P.O. Box 111, FIN-80101 Joensuu, Finland ORCID ID:E-mail:

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. https://doi.org/10.14214/sf.a9235

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, ORCID ID:E-mail:
  • Kellomäki, ORCID ID:E-mail:
  • Leinonen, ORCID ID:E-mail:
  • Repo, ORCID ID:E-mail:
article id 5430, category Article
Winai Sirikul, Markku Kanninen. (1990). Shoot growth and its clonal variation in Pinus kesiya. Silva Fennica vol. 24 no. 3 article id 5430. https://doi.org/10.14214/sf.a15584

Shoot elongation of Pinus kesiya Royle ex Gordon was studied using 2-year old grafts in a clonal seed orchard of the Pine Improvement Centre, located at the Huey Bong Experimental Station near Chiangmai, Thailand (19° 17’ N, 99° 15’ E, 900 m a.s.l.).

The seed orchard had a completely randomized block design with 30 blocks and 80 single-tree plots (clones) in each block. Eleven clones in four blocks were selected out of the total of 80 grafts (clones). From each graft, three lateral branches at the height of 1.6 m from the ground level were selected. Thus, total of 109 branches were measured. Shoot length of branches was measured between July 3, 1983 and March 11, 1984 at approximately bi-weekly intervals. Method of classical growth analysis were used in describing the shoot growth.

The annual shoot growth pattern of P. kesiya exhibited two consecutive sigmoid growth curves, i.e. it consisted of two flushes of shoot elongation, both formed by free growth. Thus, the pattern of shoot growth resembled the caribaea pattern. However, the annual shoot was composed of summer and winter shoots. These could be distinguished from each other by the reproductive organs, which always occur on winter shoot. The shoot contributed 61% of the total annual shoot length.

There were significant differences in the pattern of shoot elongation between the studied clones, which may reflect differences in the adaptation to different environmental conditions.

The PDF includes an abstract in Finnish.

  • Sirikul, ORCID ID:E-mail:
  • Kanninen, ORCID ID:E-mail:
article id 5348, category Article
Risto Häkkinen, Pertti Hari. (1988). The efficiency of time and temperature driven regulation principles in plants at the beginning of the active period. Silva Fennica vol. 22 no. 2 article id 5348. https://doi.org/10.14214/sf.a15505

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.

The PDF includes a summary in Finnish.

  • Häkkinen, ORCID ID:E-mail:
  • Hari, 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 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 7677, category Article
Risto Heikkilä, Timo Mikkonen. (1992). Effects of density of young Scots pine (Pinus sylvestris) stand on moose (Alces alces) browsing. Acta Forestalia Fennica no. 231 article id 7677. https://doi.org/10.14214/aff.7677

The study was carried out at Padasjoki, Southern Finland, where moose (Alces alces L.) density on the winter range had been over 1.5 individuals/km2. Moose browsing intensity, expressed in terms of number of twigs eaten and biomass used, increased with stand density (biomass). Total biomass consumption (dry weight) per sample plot and per sapling. The number of bites increased, but the percentage biomass removed did not differ when stand density increased. A relatively large bite size was observed on the plots of low stand density. The quantity of food, which on average was of relatively low quality, was obviously important due to the benefit gained through reducing the search time.

The nutritive value of the browse, expressed in terms of chemical compounds indicating low food digestibility, was lower in the dense than in the sparse Scots pine stand. However, the amount of crude protein and arginine were relatively high in the dense stand. We concluded that shading affected the nutritional status of saplings on high density plots.

Although the biomass removed by moose per sapling was high for low density plots, the remaining biomass was larger than that on the high-density plots owing to the relatively large twig biomass of saplings. The number of saplings per hectare without main stem breakage increased significantly as stand density increased.

The PDF includes a summary in Finnish.

  • Heikkilä, ORCID ID:E-mail:
  • Mikkonen, ORCID ID:E-mail:

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