Harri Mäkinen

Radial growth variation of Norway spruce (Picea abies (L.) Karst.) across latitudinal and altitudinal gradients in central and northern Europe

Regional and temporal growth variation of Norway spruce (Picea abies (L.) Karst.) and its dependence on air temperature and precipitation were compared in stands across latitudinal and altitudinal transects in southwestern and eastern Germany, Norway, and Finland. The temporal variation of radial growth was divided into two components: medium- and high-frequency variation, i.e. decadal and year-to-year variation, respectively. The medium-frequency component was rather different between regions, especially the southern and northern ones. However, within each region the medium-frequency growth variation was relatively similar, irrespective of altitudinal and latitudinal differences of the sample sites. A part of the high-frequency variation was common to all four regions, which suggests that some factors synchronising tree growth are common for the entire study area. The high-frequency component of growth was more strongly related to monthly air temperature and precipitation than was the medium-frequency variation. The limiting effect of low temperatures was more significant at northern as well as high-altitude sites, while the importance of precipitation increased in the south and at low altitudes.

Predicting the decomposition of Scots pine, Norway spruce, and birch stems in Finland.

Models were developed for predicting the decomposition of dead wood for the main tree species in Finland, based on data collected from long-term thinning experiments in southern and central Finland. The decomposition rates were strongly related to the number of years after tree death. In contrast to previous studies, which have used the first-order exponential model, we found that the decomposition rate was not constant. Therefore, the Gompertz and Chapman-Richards functions were fitted to the data. The slow initial decomposition period was mainly due to the fact that most dead trees remained standing as snags after their death. The initial period was followed by a period of rapid decomposition and, finally, by a period of moderately slow decomposition. Birch stems decomposed more rapidly than Scots pine and Norway spruce stems. Decomposition rates of Norway spruce stems were somewhat lower than those of Scots pine. Because the carbon concentration of decaying boles was relatively stable (about 50%) the rate of carbon loss follows that of mass loss. Models were also developed for the probability that a dead tree remains standing as a snag. During the first years after death, the probability was high. Thereafter, it decreased rapidly, the decrease being faster for birch stems than for Scots pine and Norway spruce stems. Almost all stems had fallen down within 40 years after their death. In Scots pine and Norway spruce, most snags remained hard and belonged to decay class 1. In birch, a higher proportion of snags belonged to the more advanced decay classes. The models provide a framework for predicting dead wood dynamics in managed as well as dense unthinned stands. The models can be incorporated into forest management planning systems, thereby facilitating estimates of carbon dynamics.

Woody biomass production lags stem-girth increase by over one month in coniferous forests

Wood is the main terrestrial biotic reservoir for long-term carbon sequestration(1), and its formation in trees consumes around 15% of anthropogenic carbon dioxide emissions each year(2). However, the seasonal dynamics of woody biomass production cannot be quantified from eddy covariance or satellite observations. As such, our understanding of this key carbon cycle component, and its sensitivity to climate, remains limited. Here, we present high-resolution cellular based measurements of wood formation dynamics in three coniferous forest sites in northeastern France, performed over a period of 3 years. We show that stem woody biomass production lags behind stem-girth increase by over 1 month. We also analyse more general phenological observations of xylem tissue formation in Northern Hemisphere forests and find similar time lags in boreal, temperate, subalpine and Mediterranean forests. These time lags question the extension of the equivalence between stem size increase and woody biomass production to intra-annual time scales(3, 4, 5, 6). They also suggest that these two growth processes exhibit differential sensitivities to local environmental conditions. Indeed, in the well-watered French sites the seasonal dynamics of stem-girth increase matched the photoperiod cycle, whereas those of woody biomass production closely followed the seasonal course of temperature. We suggest that forecasted changes in the annual cycle of climatic factors(7) may shift the phase timing of stem size increase and woody biomass production in the future.

Climatic signal in annual growth variation in damaged and healthy stands of Norway spruce [Picea abies (L.) Karst.] in southern Finland

Abstract. Annual radial increment variation and its dependence on temperature and precipitation were studied in 13 severely damaged and 12 healthy stands of Norway spruce [Picea abies (L.) Karst.] on mineral soil and peatlands in southern Finland. An intervention analysis revealed that the presently dead trees had experienced severe growth reductions since the late 1980s. Even though annual growth variation between the stands was fairly similar, differences in the relationship between radial growth and weather variation were found between sites. High temperature in May increased radial growth in the healthy stands. This was not observed in the damaged stands, where summer temperature was negatively correlated with growth. In addition, high temperature during the previous summer decreased tree growth in next summer. Tree-ring indices showed a strong positive correlation with June precipitation in the damaged stands. Correlation of June precipitation and growth was much weaker in the healthy stands. This result suggests that the damage is connected to drought and is likely to occur at drought-sensitive sites. The finding also fits in well with the fact that many of the damaged sites were rocky or stony.

Large-scale climatic variability and radial increment variation of Picea abies (L.) Karst. in central and northern Europe

Abstract. High-frequency variation of Norway spruce radial increment [Picea abies (L.) Karst.] and its dependence on various climatic variables was compared in stands across latitudinal and altitudinal transects in southwestern and eastern Germany, Norway, and Finland. The tested variables included local temperature and precipitation, northern hemisphere temperature anomalies, and the climatic teleconnection patterns (North Atlantic Oscillation, East Atlantic, East Atlantic Jet, East Atlantic/West Russia, and Scandinavian patterns). Climatic impact on radial increment increased towards minimum and maximum values of the long-term temperature and precipitation regimes, i.e. trees growing under average conditions respond less strongly to climatic variation. Increment variation was clearly correlated with temperature. Warm Mays promoted radial increments in all regions. If the long-term average temperature sum at a stand was below 1,200–1,300 degree days, above average summer temperature increased radial increment. In regions with more temperate climate, water availability was also a growth-limiting factor. However, in those cases where absolute precipitation sum was clearly related to radial increment variation, its effect was dependent on temperature-induced water stress. The estimated dates of initiation and cessation of growing season and growing season length were not clearly related to annual radial increment. Significant correlations were found between radial increment and climatic teleconnection indices, especially with the winter, May and August North Atlantic Oscillation indices, but it is not easy to find a physiological interpretation for these findings.

The effect of artificially induced drought on radial increment and wood properties of Norway spruce

We studied experimentally the effects of water availability on height and radial increment as well as wood density and tracheid properties of Norway spruce (Picea abies (L.) Karst.). The study was carried out in two long-term N-fertilization experiments in Southern Finland (Heinola and Sahalahti). At each site, one fertilized and one control plot was covered with an under-canopy roof preventing rainwater from reaching the soil. Two uncovered plots were monitored at each site. The drought treatment was initiated in the beginning of growing season and lasted for 60-75 days each year. The treatment was repeated for four to five consecutive years depending on the site. Altogether, 40 sample trees were harvested and discs sampled at breast height. From the discs, ring width and wood density were measured by X-ray densitometry. Tracheid properties were analysed by reflected-light microscopy and image analysis. Reduced soil water potential during the growing season decreased annual radial and height increment and had a small influence on tracheid properties and wood density. No statistically significant differences were found in the average tracheid diameter between the drought-treated and control trees. The average cell wall thickness was somewhat higher (7-10%) for the drought treatment than for the control, but the difference was statistically significant only in Sahalahti. An increased cell wall thickness was found in both early- and latewood tracheids, but the increase was much greater in latewood. In drought-treated trees, cell wall proportion within an annual ring increased, consequently increasing wood density. No interaction between the N fertilization and drought treatment was found in wood density. After the termination of the drought treatment, trees rapidly recovered from the drought stress. According to our results, severe drought due to the predicted climate change may reduce Norway spruce growth but is unlikely to result in large changes in wood properties.

Effect of Growth Rate on Fibre Characteristics in Norway Spruce (Picea abies (L.) Karst.)

Summary To study the effect of growth rate on fibre characteristics and their variations in Norway spruce, trees were sampled in a nutrient optimisation experiment in northern Sweden. Data was collected from 24 trees (40 years old) from fertilised and control plots after 12 years of annual nutrient application, as well as from older trees outside the experimental area. Fibre length, fibre diameter, cell wall thickness, lumen diameter and cell wall percentage were measured from every third annual ring at breast height and at a height of 4 m. Fibre properties, as well as their standard deviation, were closely related to ring number and distance from the pith. Intra-ring variation of fibre characteristics was high compared to their variation between trees. Fertilisation reduced fibre length and cell wall thickness, but increased fibre and lumen diameter in rings of the same age. The difference in fibre width, cell wall thickness and lumen diameter between fertilised and control trees was less apparent, but a greater difference in fibre length was found between the treatments with regard to distance from the pith. There was a similar effect of fertilisation on fibre properties in early- and latewood. The effect of enhanced growth rate was less pronounced at a height of 4 m (near the pith) than at breast height (in older rings). It was demonstrated that it is possible to model intra-tree variability of fibre characteristics using ring width and cambial age as independent variables. Models presented are, however, limited by the relatively young age of the sample trees used.

Generating 3D sawlogs with a process-based growth model

Abstract This paper reports the application of a process-based tree and stand growth model to predictions of stem structure, in particular, the 3D geometry of the stem and its internal knots, in Scots pine (Pinus sylvestris L.). The model is an extension of an earlier model which predicted tree growth and competition on the basis of carbon balance, and the vertical profile of the stem and crown on the basis of a dynamic interpretation of the pipe model theory. In this study, statistical models are used to create individual branch information from the vertical profile of branch basal area, and the resulting predictions of 3D stem structure are tested against measured trees. The model predictions of the vertical size distribution of external knots in different size classes of trees are realistic, although the model tends to underestimate knot size in the top of the crown especially in dominant trees. As regards the internal knots, the model tends to slightly underestimate the width of the tight knot zone, especially in suppressed trees. The test results suggest that the model is readily applicable to predictions of the effects of stand management on wood quality distribution. Further research topics include incorporating defects other than branch size and quality in the model, as well as testing the model against stand level quality distributions of timber.

A physiological model of softwood cambial growth

Cambial growth was modelled as a function of detailed levelled physiological processes for cell enlargement and water and sugar transport to the cambium. Cambial growth was described at the cell level where local sugar concentration and turgor pressure induce irreversible cell expansion and cell wall synthesis. It was demonstrated how transpiration and photosynthesis rates, metabolic and physiological processes and structural features of a tree mediate their effects directly on the local water and sugar status and influence cambial growth. Large trees were predicted to be less sensitive to changes in the transient water and sugar status, compared with smaller ones, as they have more water and sugar storage and were, therefore, less coupled to short-term changes in the environment. Modelling the cambial dynamics at the individual cell level turned out to be a complex task as the radial short-distance transport of water and sugars and control signals determining cell division and cessation of cell enlargement and cell wall synthesis had to be described simultaneously.

Effect of sample selection on the environmental signal derived from tree-ring series

Possibilities to improve the expressed population signal (EPS) derived from tree-ring chronology were studied on 19 trees sampled from a permanent Scots pine (Pinus sylvestris L.) thinning experiment in southern Finland. Trees of varying relative size from thinned and unthinned control plots, as well as radii in different compass directions and different positions along the stem, were compared. The EPS values increased with increasing number of chronologies sampled. The effect of increasing tree number was more pronounced compared to increasing number of radii per tree. EPS also increased when the social status of a tree improved, but no clear differences were found between unthinned and thinned plots. Furthermore, no clear trend existed between different heights along the stem. The highest EPS values occurred in the directions west, southwest, and south; correspondingly, the lowest value was on the north side of the bole, followed by the northeast and east. It was concluded that some reduction in the noise present in tree-ring chronologies can be achieved by paying more attention to sample selection within and between stems. The sunny side of the bole especially, as well as dominant trees, should be preferred. However, the most important factor in noise reduction is adequate replication.