Shoot growth and crown development: effect of crown position in three-dimensional simulations

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.     

Acclimation of photosynthetic capacity in Scots pine to the annual cycle of temperature

Coniferous trees growing in the boreal and temperate zones have a clear annual cycle of photosynthetic activity. A recent study demonstrated that the seasonal variation in photosynthetic capacity of Scots pine (Pinus sylvestris L.) could be attributed mainly to the light response curve of photosynthesis. The magnitude of the light response curve varied over the season while its shape remained constant, indicating that the two physiological parameters quantifying the curve-the quantum yield per unit internal carbon dioxide concentration and the corresponding light-saturated rate-remained proportional to each other. We now show, through modeling studies, that the quantum yield (and hence the light-saturated rate) is related to the annual cycle of temperature through a delayed dynamic response. The proposed model was tested by comparing model results with intensive measurements of photosynthesis and driving variables made from April to October in three shoots of Scots pine growing near the northern timberline. Photosynthetic capacity showed considerable acclimation during the growing season. A single model describing photosynthetic capacity as a reversible, first-order delay process driven by temperature explained most of the variation in photosynthetic capacity during the year. The proposed model is simpler but no less accurate than previous models of the annual cycle of photosynthetic capacity.     

Adaptation of the LIGNUM model for simulations of growth and light response in Jack pine

LIGNUM is a whole tree model, developed for Pinus sylvestris in Finland, that combines tree metabolism with a realistic spatial distribution of morphological parts. We hypothesize that its general concepts, which include the pipe model, functional balance, yearly carbon budget, and a set of architectural growth rules, are applicable to all trees. Adaptation of the model to Pinus banksiana, a widespread species of economic importance in North America, is demonstrated.

Conversion of the model to Jack pine entailed finding new values for 16 physiological and morphological parameters, and three growth functions. Calibration of the LIGNUM Jack pine model for open grown trees up to 15 years of age was achieved by matching crown appearance and structural parameters (height, foliage biomass, aboveground biomass) with those of real trees. A sensitivity study indicated that uncertainty in the photosynthesis and respiration parameters will primarily cause changes to the net annual carbon gain, which can be corrected through calibration of the growth rate. The effect of a decrease in light level on height, biomass, total tree branch length, and productivity were simulated and compared with field data. Additional studies yielded insight into branch pruning, carbon allocation patterns, crown structure, and carbon stress. We discuss the value of the LIGNUM model as a tool for understanding tree growth and survival dynamics in natural and managed forests.     

Migration strategies of brown trout (Salmo trutta) in a subarctic river system as revealed by stable isotope analysis

We estimated the proportions of anadromous and freshwater‐resident brown trout (Salmo trutta) in different parts of the subarctic River Näätämöjoki/Neidenelva system (Finland and Norway) using carbon, nitrogen and hydrogen stable isotope analyses of archived scales as identifiers of migration strategy. Our results showed that carbon stable isotope values were the best predictor of migration strategy. Most individuals fell into two clearly distinct groups representing anadromous (47%) or freshwater‐resident (42%) individuals, but some fish had intermediate carbon values suggesting repeated movement between freshwater and the sea. The proportion of anadromous individuals decreased steadily with distance from the sea forming a spatial continuum in migration strategies which is probably maintained by the combination of several factors such as divergent availability of food resources, variable migration costs and genetic differences. These within‐catchment differences in migration strategies should be taken into account in fisheries management practices.     

Ground vegetation has a major role in element dynamics in an ash-fertilized cut-away peatland

In Finland, recycling of wood-ash as a fertilizer has been widely studied in recent decades in peatlands drained for forestry. Wood-ash is reported to promote tree growth without any significant negative impacts on the environment and could, therefore, be a suitable option on cut-away peatlands, which are estimated to increase by 30,000 ha over the next ten years. However, the environmental effects of ash-fertilization on cut-away peatlands or on ground vegetation are not well known. We studied the effects of wood- and peat-ash application on the nutrient content in soil and plants and on the heavy metal content in the vegetation of a cut-away peatland. Six treatments of different quantities and mixtures of wood-ash, peat-ash, biotite, or Forest PK-fertilizer were replicated in three plots. Both wood- and peat-ash fertilization ensured an adequate level of nutrients for the early establishment and growth of ground vegetation and birch seedlings in a cut-away peatland. Dense pioneer moss cover was important in trapping the heavy metals, such as cadmium, which are present in the ash. Ground vegetation of mosses and herbaceous plants also had a major role in the retention of the nutrients that could otherwise have leached from the open cut-away peatland area during the early stages of afforestation. Both wood- and peat-ash proved to be suitable for the initial fertilization of afforested cut-away peatlands, but in order to guarantee a sufficient long-term potassium supply for tree growth, an application of biotite with the peat-ash is recommended.     

Status of Zoobenthos and Fish Populations in Subarctic Rivers of the Northernmost Finland: Possible Effects of Acid Emissions from Russian Kola Peninsula

In 1990, a monitoring programme was initiated to survey the status of benthic invertebrate communities and fish populations over a wide range of subarctic rivers in northernmost Finnish Lapland (68°15′–70°N). A special emphasis was placed on detecting possible effects of acidification through sulfur emissions from Russian Kola Peninsula. Sampling in 13 rivers within four major river systems covered watercourses from small brooks to large 7th order rivers, and watersheds with different distances from the emission sources. The community structure of zoobenthos and the occurrence of acid-sensitive indicator species were assessed in 90 sampling sites. Sensitive fluvial fish species, including Atlantic salmon (Salmo salar), brown trout (Salmo trutta), European minnow (Phoxinus phoxinus) and burbot (Lota lota), were monitored by studying their occurrence, abundance and juvenile recruitment. No signs of acid-induced failure in juvenile recruitment were detected in salmonid populations. The occurrence of minnow and burbot covered all the systems studied. In parallell, the structure of benthic communities with relatively high species diversity and wide distribution of a highly acid-sensitive caddisfly indicated no effects of acidification.     

Survival of reared Atlantic salmon and sea trout in relation to marine conditions of smolt year in the Baltic Sea

The marine survival of hatchery-reared Atlantic salmon (Salmo salar) and sea trout (Salmo trutta) was examined in relation to marine conditions during post-smolt migration and in relation to stock traits. In 1970–2001, Carlin-tagged smolts were released in the Iijoki and Oulujoki rivers, the northern Baltic Sea. When both species were analysed together, the abundance of the three prey fish, herring (Clupea harengus), smelt (Osmerus eperlanus) and vendace (Coregonus albula) correlated positively with the survival of salmonids. In addition, the increase in smolt size appeared to improve the survival rate. Sea surface temperature (SST) may have affected indirectly through the abundance of prey fish during the post-smolt migration of salmon and sea trout. The smelt and vendace showed a statistical effect on survival only when the temperature effects were not included in the models. In sea trout, an increasing smolt length was not significantly correlated with the survival in good herring recruitment years, but in poor years survival increased very rapidly with increasing smolt size. The recapture rates of the salmonids tended to decrease between the years 1970 and 2001. During the same time period, the June SST slightly decreased. The positive correlation between the annual summer SST and recapture rate of salmon may partly explain the decreasing trend in recapture rates. An increase in smolt size did not compensate for the decline in the recapture rate of either species.     

Use of modeled photosynthesis and decomposition to describe tree growth at the northern tree line

Growth of subarctic Scots pine (Pinus sylvestris L.) trees was investigated by a combination of process-based models and dendroecological approaches. Tree ring width indices were strongly autocorrelated and correlated with simulated photosynthetic production of the previous year and with organic matter N mineralization of the current year. An autoregressive model, with photosynthesis and N mineralization as external inputs, explained growth of the trees well. However, relationships for the period 1950-1992 differed significantly from relationships for the period 1876-1949; the slope of the regression of tree ring width index and photosynthesis was lower for the 1950-1992 period. Also, the autocorrelation structure of the data changed. First-order autocorrelation decreased and second-order autocorrelation increased from the earlier to the later period. This means that growth is becoming less sensitive to variations in photosynthetic production, whereas the relationships between growth and N mineralization are remaining fairly constant. We postulate that, although photosynthesis has increased in response to increasing CO2 concentrations, tree growth rate cannot parallel the increase in photosynthesis because potential growth rate is limited directly by temperature.