Current issue: 54(2)
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.
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.
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.
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.
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.
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.
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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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.
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.
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.