Variation in nutrient concentrations of Pinus sylvestris stems

Spatial and temporal variation of N, P, K, Ca, Mg, Mn, Zn, B and Al concentrations in Scots pine (Pinus sylvestris L.) bark and stemwood was studied in three stands at different stages of development. Concentrations of the mobile nutrients N, P and K increased in stemwood and inner bark towards the youngest tissues vertically towards the top of the stem, and horizontally from the pith to the youngest annual rings. The concentrations of Ca, Mg, Mn and Zn were greatest in the oldest tissues and decreased horizontally towards the youngest annual rings. There was a slight increase of most nutrients near the divide between sapwood and heartwood. Seasonal nutrient variation was greatest in the inner bark. The highest nutrient concentrations occurred in winter and spring. Concentrations also varied with stand age, especially in the inner bark, being lowest in the oldest stand.     

Variation in the amount and quality of litterfall in a Pinus sylvestris L. stand growing on a bog

The amount and nutrient content of the above-ground litterfall was followed for 9 years in an unfertilized, PKMgB and NPKMgB fertilized Scots pine stand growing on a drained ombrotrophic bog in eastern Finland. The annual litterfall on unfertilized plots was 1995 kg ha⁻¹, of which needles accounted for 74%. The effective temperature sum (threshold value + 5°C) explained 99% of the annual variation in the amount of needle litterfall when the data from one atypical year were excluded from the analysis. Nutrient concentrations were, except for Fe, higher in needle litter than in the other litterfall fractions. Nitrogen, P and K concentrations were low in autumn, and those of Ca and Mn high, possibly owing to variation in the mobility of elements during senescence. The annual litterfall input of N to the soil was 12.4 kg ha⁻¹, and the corresponding values for P and K were 0.08 kg ha⁻¹ and 1.81 kg ha⁻¹, respectively. Fertilization reduced needle litterfall in the first year after treatment, but had no effect thereafter. The amount of other litterfall fractions was not affected by fertilization in any of the 9 years of the study. Nitrogen, P, K and B concentrations increased in the needle litter after both fertilization treatments. The results indicate long-term cycling of fertilizer nutrients on the site.     

Changes in nutrient concentrations and nutrient release in decomposing needle litter in monocultural systems of Pinus contorta and Pinus sylvestris—a comparison and synthesis

Scots pine (Pinus sylvestris) and lodgepole pine (Pinus contorta) needle litters were compared in terms of nutrient composition and its change during decomposition. Initial nutrient composition differed between the species, with lodgepole pine needle litter having significantly higher concentrations of P, Mg and Mn. However, no difference was found for concentrations of N, Ca or K. Increases in concentrations of N, P and K during decomposition were significant in both litter types. For Ca the pattern of concentration changes followed a quadratic function as decomposition proceeded. Concentrations of Mg and Mn decreased in lodgepole pine needle litter. In Scots pine litter there was also an initial decrease, but it was followed by an increase in most incubations. For both Mg and Mn, changes in concentrations during decomposition differed significantly between species. In the late decomposition stages, concentrations of Mg and Mn became similar in both litter types. Nutrient concentrations generated by the models were compared with those of the humus (F and H) layer in the stands. The model was quite accurate in predicting concentrations of N and P for both species and the concentration of Mg for lodgepole pine. By contrast, it was not accurate in predicting concentrations of Ca and Mn. Nutrient release was estimated for the two species using both measured litterfall data and long‐term estimates, and regression models were used to predict concentration changes. Rates of release of P, Mg and Mn in the lodgepole pine stands were found to be about twice as high compared with those in Scots pine. Calcium was also released to a greater extent although the difference was not significant.     

Forest Fertilization Research in Finland: A Literature Review

This paper summarizes results from fertilization research carried out in Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] stands growing on mineral soil in Finland. The use of needle and soil analyses to indicate nutrient status and the need for fertilization are discussed along with the possibilities of increasing tree growth in various site types and stages of stand development. Doses, types of fertilizer, application time, growth increase, wood quality and the profitability of fertilization are also discussed. The stand response in terms of soil acidity, biological activity, soil fauna and resistance of trees to nutrient addition are all reviewed. The effects of fertilization on understorey vegetation, mushrooms and berries, and leaching loss of nutrients are also included.     

Nutrient retranslocation within the foliage of Pinus sylvestris

During the period 1983-1987, retranslocation of N, P, K, Ca, Mg, Mn, Zn, Fe, B and Al within the foliage of Scots pine (Pinus sylvestris L.) was studied in sapling, pole-stage and mature stands in eastern Finland. Needle concentrations of the mobile nutrients N, P, and K varied seasonally because of retranslocation. In unfertilized plots, needle contents of N, P and K decreased 62-92% during senescence and needle dry weight decreased 19-51%. In some years, needle contents of Ca, Mn, Zn, Fe, B and Al increased during senescence. Retranslocation was more efficient from needles with greater dry weight. In the fertilized, pole-stage stand, needle dry weight did not decrease at senescence four years after fertilization and nutrient content decreased less than in the other years. In the mature stand, there were no clear differences in needle dry weight and nutrient changes during senescence between years or between unfertilized and fertilized plots.     

Temporal variation in nutrient concentrations of Pinus sylvestris needles

Temporal variation in nutrient concentrations of Scots pine (Pinus sylvestris L.) needles was studied during a three‐year period in three stands of differing stages of development. Concentrations of N, P and K varied significantly between years; this variation was related to differences in needle dry weight. Concentrations of all measured nutrients (N, P, K, Ca, Mg, Mn, Cu, Zn, Fe, B) and Al varied between seasons; this variation was related to nutrient mobility and the annual physiological cycle. Concentrations of the mobile nutrients N, P and K decreased in spring and early summer during shoot and needle elongation and increased in late summer and autumn during needle senescence and litterfall. Concentrations of Mg, Cu, Zn and B followed somewhat similar patterns. The poorly mobile nutrients Ca, Mn and Fe accumulated gradually in needles during each growing season. Needle nutrient concentrations were stable during the nonactive period.     

Growth response of coniferous stands to whole‐tree harvesting in early thinnings

A series of 15 field experiments was established to quantify the growth response of first‐thinning stands of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst) to whole‐tree harvesting and to estimate the need for nutrient compensation. The experiments were undertaken in Finland, Norway and Sweden and represent a wide range of site conditions. The site index (H ₁₀₀) of Scots pine stands varied from 19 to 29 m, and that of Norway spruce stands from 28 to 36 m. Total amounts of biomass and nutrients removed were calculated based on data obtained from felled sample trees. During the first 5‐yr period the growth response to the removal of logging residues varied considerably in both pine and spruce stands. Regression analyses did not reveal any functions that explained the variation in results satisfactorily. In cases where whole‐tree harvesting influenced tree growth negatively, this effect was counteracted by compensatory fertilization. It was concluded that to determine the response of remaining trees to harvesting intensity reliably, the post‐harvest period analysed must be longer than 5 yrs.     

Response of Compensatory-Fertilized Pinus sylvestris to Infection by Heterobasidion annosum

The effects of compensatory fertilization on the infection and growth rate of Heterobasidion annosum were studied in a 40-yr-old, naturally regenerated Scots pine (Pinus sylvestris L.) stand in south-eastern Finland. The treatments were: (1) unfertilized control, (2) a compound fertilizer containing P, K, Ca, Mg, S, Cu, Zn and B, (3) as treatment 2 but with nitrogen, (4) as treatment 3 but with limestone, (5) a mixture of nitrogen and limestone composed on the basis of needle analysis, and (6) wood ash. Pine roots were inoculated on two occasions four and five growing seasons after fertilization using four different strains of the P type of H. annosum. Two years after inoculation, the fungus was alive in 28.9% of the 135 inoculated roots. Fungus survival was highest in the control trees (50%) and poorest in the treatments 2 and 3 (10.5% and 16.7%, respectively). These two treatments differed significantly from the control. Total mycelial extension during the 2-yr study period was 16.4, 2.1, 4.5, 18.1, 15.2 and 5.3 cm in treatments 1, 2, 3, 4, 5 and 6, respectively. Only treatment 2 differed significantly from the control (p<0.05, df=23). The result suggests that the application of slow-release compound fertilizers without supplementary limestone may increase the resistance of Scots pine to infection by H. annosum. The nitrogen-free compound fertilizer may also retard the spread of annosum root rot in infected pine stands.     

Impregnation of Scots pine and beech with tannin solutions: effect of viscosity and wood anatomy in wood infiltration

The impregnation process of Scots pine and beech samples with tannin solutions was investigated. The two materials involved in the process (impregnation solution and wood samples) are studied in depth. Viscosity of mimosa tannin solutions and the anatomical aspect of beech and Scots pine were analysed and correlated. The viscosity of tannin solutions presents a non-newtonian behaviour when its pH level increases, and in the case of addition of hexamine as a hardener, the crosslinking of the flavonoids turns out to be of great importance. During the impregnation of Scots pine (Pinus sylvestris L.) and beech (Fagus sylvatica L.), the liquid and solid uptakes were monitored while taking into consideration the different conditions of the impregnation process. This method allowed to identify the best conditions needed in order to get a successful preservative uptake for each wooden substrate. The penetration mechanism within the wood of both species was revealed with the aid of a microscopic analysis. Scots pine is impregnated through the tracheids in the longitudinal direction and through parenchyma rays in the radial direction, whereas in beech, the penetration occurs almost completely through longitudinal vessels.     

Models for predicting microfibril angle variation in Scots pine

Context

Microfibril angle (MFA) is one of the key determinants of solid timber performance due to its strong influence on the stiffness, strength, shrinkage properties and dimensional stability of wood.

Aims

The aim of this study was to develop a model for predicting MFA variation in plantation-grown Scots pine (Pinus sylvestris L). A specific objective was to quantify the additional influence of growth rate on the radial variation in MFA.

Methods

Twenty-three trees were sampled from four mature Scots pine stands in Scotland, UK. Pith-to-bark MFA profiles were obtained on 69 radial samples using scanning X-ray diffractometry. A nonlinear mixed-effects model based on a modified Michaelis–Menten equation was developed using cambial age and annual ring width as explanatory variables.

Results

The largest source of variation in MFA (>90 %) was within trees, while between-tree variation represented just 7 % of the total. Microfibril angle decreased rapidly near the pith before reaching stable values in later annual rings. The effect of ring width on MFA was greater at higher cambial ages.

Conclusion

A large proportion of the variation in MFA was explained by the fixed effects of cambial age and annual ring width. The final model is intended for integration into growth, yield and wood quality simulation systems.     

Modelling the distribution of wood properties along the stems of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected by silvicultural management

In this work, empirical ring-based models were developed to predict the distribution of early wood percentage, wood density and fibre length along the stems of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected by silvicultural management. The performance of the ring-based models was also compared for Scots pine and Norway spruce with corresponding disc-based (cross-sectional) models. Moreover, both models were integrated with example simulations by a process-based growth and yield model to analyze how management, such as thinning, affects the growth and wood properties of Scots pine trees over a rotation as an average for the tree stem, but also along the stem.The ring-based models built for annual early wood percentage (explained by ring width), air dry wood density (explained by early wood percentage and cambial age) and fibre length (explained by radial growth percentage and cambial age) predicted reasonably well the wood properties both at an intra-ring level, but also at a cross-sectional level. These predictions were also reasonably well in line with corresponding cross-sectional predictions by the disc-based models (which predicted the properties based on the number of annual rings and diameter at breast height and/or the cross-section being considered and temperature sum). The example simulations also demonstrated that both models predicted slightly lower wood density for dominant trees compared to dominated ones grown in thinned and unthinned Scots pine stands over a rotation. Unlike the disc-based model, the ring-based model predicted, on average, higher early wood percentage in dominant trees than in dominated ones. However, fibre length was not significantly affected when the averages of the whole stems were predicted, and this held true for both ring- and disc-based models.In summary, the incorporation of empirical ring-based wood property models into a process-based growth and yield model, offers a means to study in detail how environmental conditions, forest structure and management affect the quantity and properties of stem wood produced over a rotation. The disc-based wood property models used in this work are based on data with large geographical and genetic variation, and therefore may turn out to be more applicable for predicting future wood and fibre resources at a regional and national level. This kind of integrated use of wood property models with a process-based growth and yield model could help us to evaluate the forest resources under current and changing climate.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Impacts of bucking and delimbing alternatives on pulpwood and energy wood yields in young thinning stands in Finland

The supply of energy wood from young forests could be enhanced by altering the harvesting methods in integrated pulpwood and energy wood thinnings. In this study, effects of different bucking and delimbing options on the biomass division into energy wood, pulpwood and logging residue were estimated in integrated thinnings of young stands dominated by Scots pine (Pinus sylvestris), Norway spruce (Picea abies) or birches (Betula spp.) in Finland. Thinning options were simulated in the harvestings of model stands created on the basis of forest inventory data. Increasing the minimum top diameter of pulpwood (MTDP) expectedly increased energy wood and decreased pulpwood yields. Depending on tree species, energy wood yield increased by 36–65% when the top diameter was increased from the regular level (6 or 7 cm) to 8 cm in whole-tree harvesting, and was more than doubled when the top diameter was increased to 10 cm. Delimbed energy wood yield was increased by 240–280% when the top diameter was increased to 10 cm. Total harvesting yield in whole-tree thinnings was slightly increased by increasing top diameter. These increments in harvesting yields and reallocations of tree biomass could have an impact on large-scale forest energy potentials.     

Genetic parameters for early wood and latewood densities and development with increasing age in Scots pine

? Each annual ring in pines consists of earlywood and latewood with considerable difference in density and width. To get a better determination of the genetic regulation of total wood density in Scots pine (Pinus sylvestris L.), density and width of those ring sections were measured in annual ring numbers 12 to 21 of Scots pines in a full-sib progeny test. Tree height and stem diameter were also measured.

? Heritabilities for the annual ring sections increased with age for earlywood density from 0.08 to approximately 0.25; latewood density showed similar reductions. Heritability over all 10 annual rings was 0.25 for earlywood density, 0.22 for latewood density, 0.29 for height and 0.10 for stem diameter. Genetic correlations between earlywood and latewood density and growth traits were negative, while they were strongly positive between densities of adjacent annual rings (0.70–1.0).

? Despite the higher heritability of earlywood density, the strong positive genetic correlation between those traits indicates little benefit from focusing solely on earlywood density when selecting for wood density. Analysing earlywood and latewood separately does not benefit from including the width of the corresponding ring section as a covariate. Juvenile wood may possibly turn into mature wood 15–20 y from the pith.

     

An integrated assessment approach to optimal forest bioenergy production for young Scots pine stands

Background: Bioenergy is re-shaping opportunities and imperatives of forest management. This study demonstrates,through a case study in Scots pine(Pinus sylvestris L.), how forest bioenergy policies affect stand management strategies.Methods: Optimization studies were examined for 15 Scots pine stands of different initial stand densities, site types, and temperature sum regions in Finland. Stand development was model ed using the Pipe Qual stand simulator coupled with the simulation-optimization tool Opti For Bioenergy to assess three forest bioenergy policies on energy wood harvest from early thinnings.Results: The optimal solutions maximizing bare land value indicate that conventional forest management regimes remain optimal for sparse stands. Energy harvests occurred only when profitable, led to lower financial returns. A forest bioenergy policy which included compulsory energy wood harvesting was optimal for denser stands. At a higher interest rate(4 %), increasing energy wood price postponed energy wood harvesting. In addition, our results show that early thinning somewhat reduced wood quality for stands in fertile sites. For less fertile sites, the changes were insignificant.Conclusions: A constraint of profitable energy wood harvest is not rational. It is optimal to carry out the first thinning with a flexible forest bioenergy policy depending on stand density.     

Response of mature stands of Norway spruce (Picea abies) to boron fertilization

The effects of boron (B) fertilizer applied 10 growing seasons earlier were studied in mature Norway spruce (Picea abies (L.) Karst.) trees in long-term factorial fertilization experiments at two field sites. Needle nutrient status, above-ground and below-ground growth and δ¹³C and carbon concentrations in the annual rings were measured. Needle B concentrations varied between 4 and 19 mg kg⁻¹ on the plots that had not received B fertilization. On the B-fertilized plots they varied between 15 and 39 mg kg⁻¹. The lowest B concentrations were on the plots that had received N or NCa fertilization. Needle Mn and Zn concentrations were lower on the B plots than on the plots that had not received B fertilization, although not significantly. Mean annual volume growth was slightly higher on the B plots at the more fertile site, but not at the less fertile one. The living:dead fine root mass ratio and living fine root length were also higher on the B-fertilized plots than on the unfertilized plots, but δ¹³C was not significantly affected, suggesting that the water status of the trees was not markedly altered by the increase in root growth. The carbon concentration in the annual rings was higher in the B-fertilized trees than in the unfertilized ones, suggesting the importance of B for wood formation.     

Long‐term effects of temperature on the wood production of Pinus sylvestris L. and Picea abies (L.) Karst. In old provenance experiments

Old provenance experiments with Scots pine and Norway spruce in Finland were used for assessing the long‐term effects of the projected climatic change on forest trees. The northernmost origins showed an increase in wood production when transferred southwards into a climate with an annual mean effective temperature sum close to that which is expected in northern areas as a result of the projected climatic change. A model is constructed with the estimated changes in wood production as a function of the annual mean temperature sum at the original location and the change in the annual mean temperature sum caused by the geographical transfer. The major changes in wood production are expected to occur in the northernmost areas of tree growth.     

Quantifying the effect of pine mistletoe on the growth of Scots pine

Mistletoe infection results in substantial growth losses in mistletoe‐infected forests. This study reports and evaluates the results of retrospective analyses of radial growth of Scots pine (Pinus sylvestris) in relation to the level of infection of pine mistletoe (Viscum album ssp. austriacum). A total of 43 Scots pine trees were destructively sampled from different sites. Of these trees, 14 were uninfected and 29 were infected. Infection classes were determined using six‐class dwarf mistletoe rating system (DMRS). All needle and mistletoe biomass were removed completely and weighed for each sampled tree. Subsamples from needles and all mistletoe biomass were taken to the laboratory for oven‐dried weight determinations. Five‐cm‐thick wood discs were cut from the stem at the breast height (1.3 m) to determine annual basal area increment for the last 25 years. In addition to DMRS, new infection classes were created using mistletoe‐to‐needle biomass (MB/NB) ratio. The results showed that the radial growth losses could be as much as 41% to 64% at different infection levels. The rate of growth loss in relation to DMRS and MB/NB ratio was similar, but with a larger variability in DMRS values. The results showed that both DMRS rating and MB/NB ratio seem to be important for quantifying growth loss on Scots pine trees infected with mistletoe. The results of this study can also be invaluable in modelling the effects of mistletoe on the growth of Scots pine trees.     

帽儿山地区不同种源樟子松树轮对气候因子的响应

[目的]为了进一步了解不同种源的樟子松在帽儿山地区的生长差异,研究该地区气候因子对樟子松径向生长的影响,本文建立了来自于阿尔山、高峰、红花尔基、卡伦山、图强罕达盖和金山等7个种源樟子松树轮年表。[方法]通过年轮-气候响应分析、相关分析、主成分分析,探讨了不同种源樟子松径向生长与气候因子的响应差异。[结果]表明:7个种源地樟子松树轮年表整体变化趋势基本相似,但局部也有不同之处,除图强种源外,其余6个种源地樟子松在生长初期年轮宽度呈增加趋势。阿尔山、高峰、红花尔基和卡伦山种源地樟子松径向生长与当年2月平均温度显著正相关,罕达盖种源地樟子松径向生长与前一年4月平均温度显著正相关;高峰、图强、卡伦山种源地樟子松径向生长与当年6月降水显著正相关,阿尔山、高峰、金山、卡伦山、红花尓基、图强种源樟子松的径向生长主要与前一年8月降水显著负相关。[结论]多数种源的樟子松在初期径增长较快,与当年2月份的温度和6月份的降水量有正相关的响应性,而与前一年8月份的降水量有负相关的响应性。     

Exploratory study on the impregnation of Scots pine sapwood (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) with different hot melting waxes

Scots pine sapwood (Pinus sylvestris L.) and beech (Fagus sylvatica L.) were impregnated with five waxes. The experiments indicate deep penetration into pine sapwood. Besides the viscosity, an influence of the wax polarity is presumed. Wax penetrates pine wood deeply via the cross-section, but not sufficiently enough to impregnate longer construction elements. However, the radial wax uptake exceeds the uptake via the tangential orientation and guarantees complete soaking of the sapwood tissue. The lateral wax penetration within beech is quite low and irregular. In addition to the temperature, a prolonged process procedure is decisive for an increasing wax uptake. As such, beech wood vessel elements seem to be fully impregnable via the longitudinal surface after a longer process procedure.