Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Carbon and nitrogen release from decomposing Scots pine,Norway spruce and silver birch stumps

Stumps are the largest coarse woody debris component in managed forests, but their role in nutrient cycling is poorly understood. We studied carbon (C) and nitrogen (N) dynamics in Scots pine (Pinus sylvestris), Norway spruce (Picea abies), and silver birch (Betula pendula) stumps, which had decomposed for 0, 5, 10, 20, 30 and 40 years after clear-cutting in southern Finland. Carbon and N were released significantly faster from birch stumps than from conifer stumps. In 40 years, conifer stumps lost 78% and birch stumps 90% of their initial C. In contrast, the amount of N in stumps increased, indicating that external N accumulated in the stumps. After 40 years of decomposition, the amount of N was 1.7 and 2.7 times higher than the initial amount in pine and spruce stumps, respectively. Nitrogen was released from birch stumps, but only after they had decomposed for 20 or more years. On average, 59% of N stored in birch stumps was released during 40 years. The results indicate that the stumps of the major tree species in Fennoscandian forests are long-term C and, especially, N pools which serve as N sinks, thus potentially diminishing N leaching into ground water and watercourses after harvesting. This suggests that the removal of stumps for bioenergy production may markedly affect the nutrient status and nutrient cycling of boreal forests.     

Influence of different nursery container media on rooting of Scots pine and silver birch seedlings after transplanting

Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth.) seedlings were grown in containers filled with growth media based on medium-textured sphagnum peat, coarse perlite and fine sand. The seedlings were then planted into fine and coarse sandy soils in 2.2 l pots, which were subjected to two water-content treatments (only one for birch). After the seedlings had grown five weeks in a greenhouse, rooting into the surrounding soil and shoot growth were measured. Addition of perlite and sand to peat medium slightly affected rooting; thus suggesting minor effects on seedling establishment. However, nitrogen concentration of the seedlings varied between growth media and correlated positively with rooting into the soil. The particle size and water content of the soil affected considerably rooting of the seedlings. Seedling height at the time of planting did not affect rooting or shoot growth. The fact that the fewest out-grown roots occurred in the dry fine sandy soil, suggests that dry soil together with high strength and resistance to root penetration reduce rooting and water uptake by container seedlings most and may thus cause water and nutrient stresses to seedlings after outplanting.     

Phosphorus and base cation accumulation and release patterns in decomposing Scots pine,Norway spruce and silver birch stumps

Tree stumps are integral constituents of managed forest ecosystems, but their role in nutrient cycling is poorly understood. We studied phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) dynamics in decomposing Scots pine (Pinus sylvestris), Norway spruce (Picea abies), and silver birch (Betula pendula) stumps in southern Finland in a chronosequence of 0-, 5-, 10-, 20-, 30- and 40-year-old clear-cut areas. Along with the decomposition of pine and spruce stumps, the amount of P in stumps increased, but K and Ca were released, and the amount of Mg initially decreased and then increased. All nutrients, except K, accumulated in birch stumps during the first ten years, but were released thereafter. After 40 years of decomposition, pine and spruce stumps contained 180% and 202% of their initial amounts of P, respectively. In addition, the amounts of Mg were larger than the initial amounts in 40-year decomposed pine (126%) and spruce (202%) stumps. In contrast, birch stumps lost 64% and 75%, respectively, of their initial amounts of P and Mg over a 40-year period. The stumps of all the species were found to release K and Ca. Pine, spruce and birch stumps lost 48%, 64% and 87% of their initial amount of K, and 49%, 35% and 42% of their initial amount of Ca, respectively, during the 40-year period. The results indicate that decomposing stumps of the major tree species in Fennoscandian forests are long-term nutrient pools and they serve as P sinks, thus potentially reducing P leaching after clear-cutting.     

Dissolved organic carbon and nitrogen leaching from Scots pine, Norway spruce and silver birch stands in southern Sweden

The effects of three common tree species - Scots pine, Norway spruce and silver birch - on leaching of dissolved organic carbon and dissolved nitrogen were studied in an experimental forest with podzolised soils in southern Sweden. We analyzed soil water collected with lysimeters and modeled water fluxes to estimate dissolved C and N fluxes. Specific UV absorbance (SUVA) was analyzed to get information about the quality of dissolved organic matter leached from the different stands. Under the O horizon, DOC concentrations and fluxes in the birch stands were lower than in the spruce and pine stands; annual fluxes were 21 g m⁻² y⁻¹ for birch and 38 g m⁻² y⁻¹ and 37 g C m⁻² y⁻¹ for spruce and pine, respectively. Under the B horizon, annual fluxes for all tree species ranged between 3 and 5 g C m⁻² y⁻¹, implying greater loss of DOC in the mineral soil in the coniferous stands than in the birch stands. We did not find any effect of tree species on the quality of the dissolved organic matter, as measured by SUVA, indicating that the chemical composition of the organic matter was similar in leachates from all three tree species. Substantial amounts of nitrogen was leached out of the soil profile at the bottom of the B horizon from the pine and birch stands, whereas the spruce stands seemed to retain most of the nitrogen in the soil. These differences in N leaching have implications for soil N budgets.     

Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

The effects of soil and air temperature on CO2 exchange and net biomass accumulation in Norway spruce, Scots pine and silver birch seedlings

Soil temperature is proposed to affect the photosynthetic rate and carbon allocation in boreal trees through sink limitation. The aim of this study was to investigate the effect of temperature on CO(2) exchange, biomass partitioning and ectomycorrhizal (ECM) fungi of boreal tree species. We measured carbon allocation, above- and below-ground CO(2) exchange and the species composition of associated ECM fungi in the rhizosphere of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies K.) and silver birch (Betula pendula Roth) seedlings grown in soil maintained at 7-12, 12-15 and 16-22 °C. We found increased root biomass and photosynthetic rate at higher soil temperatures, but simultaneously with photosynthesis rate, higher temperature generally increased soil respiration as well as shoot, and root and rhizosphere respiration. The net CO(2) exchange and seedling biomass did not increase significantly with increasing temperature due to a concomitant increase in carbon assimilation and respiration rates. The 2-month-long growth period in different soil temperatures did not alter the ECM fungi species composition and the below-ground carbon sink strength did not seem to be directly related to ECM biomass and species composition in any of the tree species. Ectomycorrhizal species composition and number of mycorrhiza did not explain the CO(2) exchange results at different temperatures.     

Identification of multiple Endocronartium pini infections in Scots pine with PCR

Rust susceptibility in pine is known to be heritable, which means that the disease risk remains in a stand. Also, if the climate becomes warmer and more humid, as expected, the risk of rust disease will increase. The main trait of interest and importance in an obligate parasite, such as Endocronartium pini is the variation in pathogenicity. However, to test the variation in pathogenicity the fungal population in a stand must be described. In this study it was possible to partly describe an E. pini population at stand level with RAPD-PCR and identify multiple infections in a single infected tree (Pinus sylvestris). Some RAPD phenotypes detected were more common than others. This suggests some mechanism by which some phenotypes spread more effectively. The E. pini population structures in the two Pinus sylvestris stands studied were also significantly different, even when the stands shared some equally common RAPD phenotypes. The most common phenotype was not always present in a tree with multiple infections, as would be expected. Thus, some specific resistance interaction may occur. Further research is needed to prove this assumption.     

Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland

Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.     

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.     

Boron retranslocation in Scots pine and Norway spruce

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.     

Ectomycorrhizal communities associated with silver fir seedlings (Abies alba Mill.) differ largely in mature silver fir stands and in Scots pine forecrops

Context

The requirement for rebuilding forecrop stands besides replacement of meadow vegetation with forest plants and formation of soil humus is the presence of a compatible ectomycorrhizal (ECM) fungal community.

Aims

This study aims to assess ectomycorrhizal fungi diversity associated with silver fir (Abies alba Mill.) seedlings regenerating in silver fir stands and Scots pine forecrops.

Methods

One-year-old seedlings were sampled in six study sites: three mature fir forests and three pine forests. ECM fungi were identified by polymerase chain reaction amplification and sequencing of the internal transcribed spacer of rDNA.

Results

The mean mycorrhizal colonization exceeded 90 %. Thirty-six ectomycorrhizal taxa were identified in fir stands and 23 in pine forecrops; ten out of these species were common to both stands. The fungal communities were different between study sites (R?=?0.1721, p?=?0.0001). Tomentella stuposa was the only species present at all sites.

Conclusion

Silver fir seedlings in Scots pine forecrops supported smaller ECM fungal communities than communities identified in mature silver fir stands. Nevertheless, fungal colonization of seedling roots was similar in both cases. This suggests that pine stands afforested on formerly arable land bear enough ECM species to allow survival and growth of silver fir seedlings.     

Estimation of nutrient removals in stem-only and whole-tree harvesting of Scots pine, Norway spruce, and birch stands with generalized nutrient equations

Whole-tree harvesting (WTH), where logging residues are removed in addition to stems, is widely practised in Fennoscandian boreal forests. WTH increases the export of nutrients from forest ecosystems. The extent of nutrient removals may depend on tree species, harvesting method, and the intensity of harvesting. We developed generalized nutrient equations for Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies Karsten), and birch (Betula pendula Roth and Betula pubescens Ehrh.) stands to be able to calculate the amounts of nitrogen, phosphorus, potassium, and calcium in stems and above-ground biomass (stem and crown) as a function of stand volume. The equations were based on Fennoscandian literature data from 34 pine, 26 spruce, and 5 birch stands, and they explained, depending on the tree species and nutrient, 61–99% and 56–87% of the variation in the nutrient amounts of stems and above-ground biomass, respectively. The calculations based on the equations showed that nutrient removals caused by stem-only harvesting (SOH) and WTH per harvested stem m³ were smaller in pine than in spruce and birch stands. If the same volume of stem is harvested, nutrient removals are, in general, nearly equal at thinnings and final cuttings in SOH, but larger in thinnings than final cuttings in WTH. If the principal aim is to minimize the nutrient removals per harvested stem m³, the harvesting should be done at mature pine stands. The effect of biomass removal on overall site nutrient status depends on site-specific factors such as atmospheric deposition, weathering of minerals, and the size of the nutrient pools in the soil.     

Needle polyamine concentrations and potassium nutrition in Scots pine

The response of free polyamines (putrescine, spermidine and spermine) in needles of Scots pine (Pinus sylvestris L.) to varying needle potassium concentrations was investigated in two potassium fertilization experiments on drained peatlands. A significant negative correlation was observed between putrescine and potassium concentrations in needles. Putrescine responded more sensitively to decreasing needle K concentrations during the growing season than during the winter. Putrescine accumulation started when needle potassium concentrations were above 5.5 mg g(dw) (-1) in summer and above 5.0 mg g(dw) (-1) in winter. A decrease in needle potassium concentrations below 4.3 mg g(dw) (-1) in summer and below 3.5 mg g(dw) (-1) in winter resulted in an exponential increase in putrescine concentrations. Putrescine accumulation was initiated well above the needle potassium concentrations generally considered indicative of deficiency. The exponential increase in putrescine concentrations in winter occurred within the range of needle potassium concentrations that has been identified as indicating severe potassium deficiency.     

Influence of nurse birch and Scots pine seedlings on early aerial development of European beech seedlings in an open-field plantation of Central France

Beech woodland can be restored by direct planting of beech (Fagussylvatica L.) seedlings in abandoned areas, but this methodis generally avoided in forestry because of the growth difficultiesof beech in full-light conditions. This study tested a methodthat consists of planting beech seedlings in full-light conditionswith silver birch (Betula pendula Roth.) or Scots pine (Pinussylvestris L.) as nurse trees. A total of 65 two-year-old beechseedlings surrounded by either 3, 4, 5 or 6 pine or birch seedlingsor without competitors were planted in bare-soil open-fieldconditions in Central France. Tree growth and light availabilitywere monitored at the end of each of the following 3 years,and soil moisture was also measured the last year. At the endof the experiment, aerial biomass measurements were made onthe beeches. Results showed that relative beech growth in termsof diameter was significantly reduced by the local neighbouringtrees and that this reduction was particularly noticeable inthe pine treatments where there was extensive pine seedlingdevelopment. In contrast, relative beech height did not exhibitany significant variations among the treatments. Light availabilitywas decreased by the neighbouring trees, especially in the pinetreatments. Light reduction was more pronounced in the last2 years and in the middle or lower parts of the canopy of thesubject beech. Soil water content was lower under the pine canopiesthan under the birch canopies, and was positively correlatedto beech relative diameter growth. Specific leaf area, height-to-diameterratio and crown length-to-crown width ratio weakly but significantlyincreased with competition from the neighbours. Beech biomasspartitioning was only weakly affected by the treatments. Thisstudy showed that nurse trees tend to reduce beech growth butimprove form although effects remained weak due to the insufficientlength of the experiment. Further studies are also needed toquantify the effects of the neighbourhood on the growth of competitiveherbaceous vegetation and on changes in the microclimatic conditions.     

Stand Density Control Diagrams for Scots pine and Austrian black pine plantations in Bulgaria

Stand Density Control Diagram (SDCD) is a stand-level mathematical model, which describes the relationships between yield, density and mortality throughout all stages of stand development. The SDCD is primarily used to derive density control schedules by management objectives. The main objectives of the present study are to define a modified model of SDCD for application to Scots pine (Pinus sylvestris L.) and Austrian black pine (Pinus nigra Arn.) plantations in Bulgaria, to examine the fitness of the model with representative experimental data sets from plantations of both species and to present a way of direct application of the SDCDs for practical purposes. The constructed SDCDs characterize the spatial-temporal dynamics of the pine plantations in a broad range of densities, forest sites and growth stages from 4 to 26 (28) m of dominant height class. The full density lines were fixed with self-thinning exponents α = 1.69 and α = 1.75 for Scots pine and Austrian black pine, respectively, and the trajectories of natural thinning for 23 initial densities (444-40000/ha) were determined. A direct way for application of the SDCDs to the plantation management was designed to estimate the optimal initial densities for the maximum attainable final yield and large-size wood production by self-thinning stands.     

Basal sweep and compression wood in young Scots pine trees

This study demonstrates the correlation between stem form and compression wood content in 36 sampled trees from a 6-yr-old container grown Scots pine ( Pinus sylvestris L.) plantation in central Sweden. Root analyses were carried out to examine causes for the high incidences of basal sweep. On the sampled trees basal sweep, stem sweep and maximum bow-height were measured. Discs were cut at different heights and the compression wood content was analysed using digital image analysis. Pith eccentricity and out-of-roundness were calculated for each disc as well as per stem. Compression wood and pith eccentricity was most pronounced near the stem base, where sweep tends to be high. In general, pith eccentricity increased with degree of basal sweep. As a conclusion, expression of basal sweep can be used to predict compression wood content in young Scots pine trees. This study also shows the importance of anchoring of a tree to prevent basal sweep formation due to instability.     

Thinning intensity and growth response in SW-European Scots pine stands

The effect of different thinning intensities on growth and yield was studied in Pinus sylvestris L. stands at the south-western limit of its distribution area (Central Spain), using five long-term thinning trials. Data were analysed collectively considering several factors (trial, block, plot and period) as random effects. Total volume and volume increment decreased with thinning intensity, this loss being more significant in the case of moderate and heavy thinning. No difference was found among treatments for total basal area or the increment in basal area. The results revealed an optimum basal area (Assmann’s definition) between 85 and 100% of the basal area in unthinned plots. Volume growth loss associated with heavy thinnings (reduction of 18% in volume increment) was smaller than that reported in Central and Northern European regions (greater than 25%). Height increment was not influenced by thinning, whereas dominant and quadratic mean diameter increments increased with the thinning intensity. The response of diameter growth to thinning was greater at younger ages (less than 50 y) and in medium-sized trees.     

Comparative analyses on the litter in Korean pine Mongolian Scots pine and Dahurian Larch plantations

Comparative analyses were conducted on the nutrient element content and returning amount of main fractional compositions of litter in Korean pine (KP), Mongolian Scots pine (MSP) and Dahurian larch (DL) plantations in Laoshan Plantation Experiment Station of Maoershan Experiment Forest Farm of Northeast Forestry University. The results are as follows: (1) The nutrient element content and returning amount in litter varies among different fractional compositions and tree species, the total returning amount of all nutrient elements and the returning amount of K, Ca, Mg, N and P are DL > MSP > KP, the returning amount of Cu is DL > KP > MSP, the returning amount of Fe and Mn are MSP > DL > KP; (2) To KP and DL plantations, the main nutrient element returned is dead needles; dead branches, bark scales and dead cones account for a little proportion; whereas to MSP plantation, besides dead needles, dead branches and bark scales also play an important role in the return of nutrient elements; (3) A little deal of dead leaves can provided a great deal of returning amount of nutrient elements.