Inoculation of young pine seedlings with Peridermium pini from north-east Scotland

Seedlings of Pinus sylvestris, P. nigra ssp. laricio, and P. mugo vars mughus, rostrata and pumilio were inoculated at their cotyledon stage with aeciospores of Peridermium pini from north-east Scotland. Infection was determined by the presence of both mycelia and haustoria in free-hand or paraffin sections. In the second growing season, infected seedlings showed symptoms such as stem-swelling, development of spermogonia and aecia, and death. In a second experiment, young P. sylvestris seedlings from seven seed sources collected in Great Britain were inoculated and the infection were examined after 6 weeks. In the samples of 50 seedlings from each source, 30–70% seedlings were infected.     

Growth patterns from different forest generations of Scots pine in Estonia

There is strong evidence that climate change alters tree growth in boreal forests. In Estonia, the analysis of ring measurements has been a common method to study growth changes. In this study, annual height growth data from dominant or co-dominant Scots pine (Pinus sylvestris L.) trees were used to develop a growth model for three forest generations. Stem analysis was applied and annual height growth was measured as the distance between whorls, as detected by branch knots of whorls on the split stem surface. Retrospective analysis of height growth produced comparative trends for three different age groups. Statistical analyses were used to estimate the impact of different factors on growth. Annual height growth was considered the best indicator for detecting possible trends in the growth potential of trees. Study results indicate that three generations (separated by time periods of 30–40 years) showed significant differences in growth patterns caused by shifts in climatic factors and management regimes (anthropogenic and natural disturbances).     

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.     

Above- and belowground biomass and net primary production in a 73-year-old Scots pine forest

We estimated above- and belowground biomass and net primary production (NPP) of a 73-year-old Scots pine (Pinus sylvestris L.) forest stand in the Belgian Campine region. Total biomass for the stand was 176 Mg ha(-1), of which 74.4% was found in stems. The root system contained 12.6% of total biomass, most of it in coarse roots (> 5 mm). Fine roots (< 5 mm) comprised only about 1.7% of total biomass, and more than 50% of fine root biomass was retrieved in the litter layer and the upper 15 cm of the mineral soil. The ratio of belowground biomass to aboveground biomass was 0.14, which is lower than that of other Scots pine forests and other coniferous forests. Between 1995 and 2001, mean annual NPP was 11.2 Mg ha(-1) year(-1), of which 68.7% was allocated to aboveground compartments. Stems, needles and cones made relatively high contributions to total NPP compared with branches. However, branch NPP was possibly underestimated because litterfall of big branches was neglected. The proportion of total NPP in belowground components was 31.3%. Coarse root NPP (2% of total) was low compared with its biomass. Fine root NPP was 3.3 Mg ha(-1) year(-1), representing about 29.5% of total NPP; however, the estimate of fine root NPP is much more uncertain than NPP of aboveground compartments. The ratio NPP/GPP (gross primary production) was 0.32, which was low compared with other coniferous forests.     

Spatial structure of semi-natural and plantation stands of Scots pine (Pinus sylvestris L.) in northern Scotland

The success of current initiatives to maintain and enhance thearea of and the special habitats provided by the remnant semi-naturalpinewoods of northern Scotland will depend upon foresters' abilityto foster more natural structures in even-aged plantations throughstand manipulation. However, there is little information onthe structures and spatial patterns that can be found in Scottishpinewoods; such knowledge could be used to design appropriatesilvicultural regimes. A study was carried out to compare spatialstructure in three 0.8–1.0 ha plots in the CairngormsNational Park; one plot was a 78-year-old plantation stand,the other two were semi-natural stands with trees up to 300years old. Basic mensurational data showed that the semi-naturalstands were characterized by a wider range of tree sizes andmore large (>50 cm d.b.h.) trees. Spatial structure was evaluatedwith a range of different indices: the aggregation index ofClark–Evans (CE), the uniform angle and diameter differentiationindices, Ripley's L function of tree spatial distribution, pairand mark correlation functions and experimental variograms oftree diameter. The CE revealed a regular distribution in theplantation with the semi-natural stands having a random pattern.Further analysis of the latter stands indicated that, in eachcase, the older trees in the stand were regularly distributedwhile the younger ones were clustered. There was little differencein uniform angle values between the stands while the diameterdifferentiation distributions suggested greater variety in diameterwithin the semi-natural stands than in the plantation. The Ripley'sL function showed that trees in the plantation were regularlydistributed at close distances but clustered over wider distances.There were differences in pattern between the semi-natural stands;in one, trees were clustered because the positions of the youngertrees were influenced by past regeneration trials, whereas inthe other stand a random pattern was observed. Similarly, thevariogram indicated widespread homogeneity in diameter withinthe plantation, while the semi-natural stands showed high variationat close spacing because of competition followed by spatialautocorrelation up to 20 m distance. Thereafter, one of thesestands had a very different pattern because of a more intensiveregeneration history. All the indices, apart from uniform angle,were able to discriminate between the plantation and the twosemi-natural stands, but only the more detailed spatial indiceswere capable of identifying differences within the latter. Theimplications of these results for management strategies in plantationsare discussed.     

The effect of brown-rot decay on water adsorption and chemical composition of Scots pine heartwood

? The effect of brown-rot (Coniophora puteana) decay on the water adsorption capacity and concentration of extractives of Scots pine (Pinus sylvestris L.) heartwood were studied by comparing corresponding properties of decayed and undecayed wood samples.

? The samples derived from 39 felled trees having a large between-tree variation in the extractive concentrations, and subsequently in the mass loss in the decay test. The water adsorption capacity, expressed as equilibrium moisture content (EMC), was measured at a high relative humidity (RH ～100%, 21 °C).

? In contrast to the widely held belief, the water adsorption capacity of brow-rotted heartwood appeared to be significantly higher than that of undecayed heartwood.

? The chemical composition of heartwood was changed radically by the fungus: the concentration of stilbenes, resin acids and free fatty acids decreased, while the concentration of soluble sugars increased as a result of decay. In addition, fungal sugars were found in the decayed samples. The concentration of total phenolics increased, which obviously reflected chemical changes in cell wall constituents other than extractives.

? As a conclusion, the information concerning the hygroscopicity of brown-rotted wood might be valuable e.g. when carrying out repairs on buildings damaged by advanced decay.

     

Stand structure and dynamics of four native Scots pine (Pinus sylvestris L.) woodlands in northern Scotland

Stands in four native pinewoods (Glenmore, Black Wood of Rannoch,Glen Garry and Glen Affric) dominated by Scots pine (Pinus sylvestrisL.) with contrasting management histories and climates wereassessed for differences in age, structure and dynamics. Treeage, height, diameter at breast height (d.b.h.), basal area,stem density and x, y coordinates were used to compare the recruitmentof trees (>1.3 m height, >7 cm d.b.h.), saplings (>1.3m height, <7 cm d.b.h.) and seedlings (<1.3 m) followingdisturbance events and protection from browsing. There was a20-year lag between localized intensive cultivation and treerecruitment on sites that were protected from deer browsing(Glenmore and Glen Garry). Recruitment was low in sites withdisturbance but no protection (Black Wood of Rannoch). The oldestpopulation, Glen Affric, showed signs of initial intense recruitmentfollowed by a long period of nil recruitment. Abundant standingdead trees were recorded only in Glen Affric, and prolific birchand rowan only in Glen Garry. Managers should consider localizedintense cultivation in conjunction with a complete reductionin browsing pressure for rapid seedling recruitment and increasedstructural diversity.     

Effect of heavy metal pollution of forest soil on radial growth of Scots pine

A field experiment with artificial soil contamination by the heavy metals copper and nickel was established in Kola Peninsula, northern Russia. To study the effect of soil contamination by heavy metals without SO₂ air pollution on the Scots pine trees growth the experiment was located outside the range of forest damage due to emissions from a copper–nickel smelter. Retardation in tree diameter growth over 15 years after contamination with 20 kg of metallurgical dust per 100 m² plot was 54%, 10 kg 27%, 5 kg 16% and 2.5 kg 7%, respectively, compared with growth in control plots. Thus, decreasing Scots pine growth due to heavy metal pollution was demonstrated.     

Relating sulfate adsorption to soil properties in Japanese forest soils

The relationship of sulfate adsorption capacity to the presence of aluminum, iron oxides, and organic carbon in four Japanese forest soils was investigated. Sulfate adsorption capacities were higher in subsoils than in surface soils, and their profiles were the inverse of those of organic carbon content. The black soil derived from volcanic ash had high sulfate adsorption capacity and showed no release of sulfate, while some of the surface horizons of the other soils desorbed sulfate. Oxalate extractable aluminum (Al₀) and iron (Fe₀) and ditionite-citrate-bicarbonate (DCB) extractable iron (Fe_d) were positively correlated with the sulfate adsorption capacity, and their correlation coefficients were almost identical (0.58–0.61). The lack of significant correlation between Al_d and the adsorption capacity suggests that DCB is unsuitable for extracting aluminum compounds which have the ability to adsorb sulfate. In contrast with oxides, organic carbon correlated with sulfate adsorption capacity negatively, and had a negative coefficient in a multiple regression equation, insignificantly. Furthermore, H₂O₂-treated samples containing decomposed organic matter adsorbed more sulfate than untreated samples. These results clearly indicated that organic matter hinders sulfate adsorption. A comparison of theβ value (standard regression coefficient) of organic carbon with that of Al₀ showed that the negative contribution of organic matter to sulfate adsorption capacity was slightly less than the positive contribution of oxides.     

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.     

Control of heather (Calluna vulgaris (L.) Hull) by steam treatment: Effects on establishment and early growth of Scots pine

Following boreal forest ecosystem disturbance,such as fire and clear-cutting, the ericaceousspecies Calluna vulgaris often becomescompletely dominant and may convert forestlandinto heathland. Calluna is known toeffectively exclude other vegetation and causes``growth check' or stagnation of coniferseedlings to result in poor tree regeneration.In this study we investigated the use of steamtreatment as an alternative method forvegetation control of Calluna vulgaris.Scots pine (Pinus sylvestris L.)establishment and growth were recorded in botha planting and a seeding experiment. Steamtreatment was compared with mechanical soilscarification and intact vegetation in theplanting experiment and with intact vegetationin the seeding experiment. The results showthat the vegetation is still strongly reducedfive years after the treatment. The steamtreatment strongly favoured the establishmentand growth of both seeded and planted Scotspine. The planted seedlings grown in the steamtreated plots had a dry weight more than twiceas high when compared to scarified plots.Seedling height and basal diameter were alsohighest for seedlings grown in steamed plots.Seedlings planted in scarified plots did notgrow as well during the first years possiblydue to injuries caused by frost heaving. Theseeding experiment showed an increasedestablishment and growth of Scots pineseedlings in steam treated plots compared to inintact vegetation.     

Modeling individual tree height growth of Norway spruce and Scots pine from national forest inventory data in Norway

We developed individual tree height growth models for Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Norway based on national forest inventory data. Potential height growth is based on existing dominant height growth models and reduced due to competition by functions developed in this study. Three spatially explicit and two spatially non-explicit competition indices were tested. Distance effects and diameter ratio effects were estimated from the data simultaneously with parameters of the potential modifier functions. Large height measurement errors in the national forest inventory data caused large residual variation of the models. However, the effects of competition on height growth were significant and plausible. The potential modifier functions show that height growth of dominant trees is largely unaffected by competition. Only at higher levels of competition, height growth is reduced as a consequence of competition. However, Scots pine also reduced height growth at very low levels of competition. Distance effects in the spatially explicit competition indices indicated that the closest neighbors are most important for height growth. However, for Scots pine also competitors at larger distance affected height growth. The five competition indices tested in this study explained similar proportions of the variation in relative height growth. Given that unbiased predictions can only be expected for the same plot size, we recommend a spatially explicit index, which describes the distance function with a negative exponential, for use in growth simulators.     

Low- to moderate-severity historical fires promoted high tree growth in a boreal Scots pine forest of Norway

Abstract

Fire is the most important ecological factor governing boreal forest stand dynamics. In low- to moderate-severity fires, the post-fire growth of the surviving trees varies according to fire frequency, intensity and site factors. Little is known about the growth responses of Scots pine (Pinus sylvestris L.) following fires in boreal forests. We quantified changes in tree growth in the years following 61 historical forest fires (between 1210 and 1866) in tree-ring series collected from fire-scarred Scots pine trees, snags and stumps in Trillemarka nature reserve in south-central Norway. Basal area increment 10 years pre-, 5 years post-, and 11–20 years post-fire were calculated for 439 fire scars in 225 wood samples. We found a slight temporary growth reduction 5 years post-fire followed by a marked growth increase 11–20 years post-fire. Beyond 20 years post-fire, the long-term tree growth declined steadily up to approximately 120 years. Our results indicate that recurring fires maintained high tree growth in remnant Scots pines, most probably due to a reduction in tree density and thus decreased competition.     

Photosynthetic utilization efficiency of absorbed photosynthetically active radiation by Scots pine and birch forest stands in the southern Taiga

Absorption and utilization of photosynthetically active radiation (PAR) were investigated in Scots pine (Pinus sylvestris L.) and birch (Betula pendula Roth.) stands that were 41 years old at the end of the experimental period. Canopy depth of the Scots pine stand was about half that of the birch stand (6.5 versus 11.0 m), but absorption of PAR was similar in the two stands. The Scots pine forest canopy, with a leaf area index of 8.9, absorbed 90% of the incoming PAR (APAR), whereas the birch forest canopy, with a leaf area index of 5.9, absorbed 92% of APAR. During maximum foliage development, the upper Scots pine canopy absorbed more PAR than the upper birch canopy (75 versus 66%). The upper, middle and lower layers of the Scots pine canopy contained 37, 48 and 15% of the total needle surface area, respectively. The corresponding distribution of foliage surface area in the three layers of the birch canopy was 50, 30 and 20%, respectively. Measurements of photosynthetic rate were combined with estimates of leaf area index and stand phytomass to determine rates of primary production on a sunny day, a cloudy day, and on an annual basis. The energy equivalents of short- and long-term carbon gain were used with determinations of APAR to calculate photosynthetic utilization efficiency. Throughout the growing season, photosynthetic utilization efficiency of APAR in the upper canopy layer of the Scots pine forest was almost twice that in the lower canopy layer. In the birch forest, photosynthetic utilization efficiency was greater in the lower canopy layer than in the upper canopy layer. In all cases, utilization efficiency was higher in the birch stand than in the Scots pine stand (52 versus 29 J kJ(-1)). Taking account of respiration of the non-photosynthetic parts of each stand (night respiration of needles or leaves; respiration of branches, trunk and roots), estimated utilization efficiency of APAR for net primary production was 11 J kJ(-1) for Scots pine and 19 J kJ(-1) for birch. Solar conversion ratios, expressed as whole-plant net primary productivity per unit of APAR for the growing season, were 0.81 g MJ(-1) for Scots pine and 0.93 g MJ(-1) for birch.     

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.     

Recovery of photosynthetic capacity in Scots pine: a model analysis of forest plots with contrasting soil temperature

Both aboveground and belowground climate affects net primary production (NNP) and forest growth. Little is known about how above and belowground factors interact. The BIOMASS-model was tested to simulate photosynthetic recovery over a wide range of soil temperatures created by snow cover manipulations on tree-scale plots in a 20-year-old Scots pine stand in northern Sweden. The differences in timing of soil warming between the plots covered a span of two months. Carbon assimilation in needles, sap flow, needle water potential and climatic parameters were measured in the field. The simulations revealed that an early start of soil warming gave a relatively early photosynthetic recovery and a 7.5% increase of NPP. Late soil warming delayed the photosynthetic recovery and reduced the NPP by 13.7%. This indicated that soil temperature needed to be accounted for, as well as air temperature, when analysing photosynthetic recovery and NPP in boreal environment. The effects of differences in soil temperature were reflected in the simulated photosynthetic recovery. The model did not fully capture the delay of photosynthetic recovery caused by a late soil warming. It was possible to integrate the complexity of the soil climate effects into a threshold date for soil thaw, using sapflow measurements together with information about air temperature and a day degree sum, as long as water availability was not limiting water uptake by roots. Although a more realistic mechanism than that currently in BIOMASS is desirable as climate change shifts the typical patterns of interplay between air and soil temperature dynamics.     

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.     

Differences in Agrobacterium infections in silver birch and Scots pine

The aim of this study was to investigate the differences in infections caused by Agrobacterium tumefaciens in a conifer, Scots pine (Pinus sylvestris), and in a non-host deciduous species, silver birch (Betula pendula). All the Agrobacterium tumefaciens strains tested caused crown-gall formation in both tree species, but the infection rates varied remarkably. In Scots pine, the development of galls was rare, and slower than in silver birch. Inoculation into the base of the stem were the most successful in gall induction. Silver-birch galls were large, often surrounding the whole stem, in contrast to Scots pine galls, which were characterized by their small size and neck-like connection with the host plant. In silver birch, no other morphological changes could be seen. In Scots pine, abnormal phenotypes with proliferating short shoots above the galls were observed during the second and third growing season. The results indicate that, of the two non-host tree species, the deciduous one, silver birch, is more susceptible to an A. tumefaciens infection than the conifer, Scots pine. The matrix for A. tumefaciens infection in silver birch differs from that in Scots pine, since the terpene compounds of Scots pine seem either to kill the agrobacteria or to suppress their growth. The differences between the species could be partly caused by their difference in sensitivity to phytohormones. These features reflect evolutionary incompatibility between A. tumefaciens and a gymnosperm.     

Acid phosphatase activity and phosphorus nutrition in Scots pine needles

From March to October, acid phosphatase activity and phosphorus (P) concentration were measured in four needle age classes of the upper and lower crowns of fertilized and unfertilized Scots pine (Pinus sylvestris L.) trees. Negative correlations between acid phosphatase activity and P concentration were observed in current-year needles and in needles in the upper part of the crown, whereas there was a positive correlation between enzyme activity and P concentration in older needles and in needles in the lower part of the crown. In May and October, needles of all ages showed increased acid phosphatase activity. The most sensitive response of acid phosphatase activity to phosphate supply and phosphorus status of the whole tree was seen in current-year needles on the first whorl where a 300% increase in acid phosphatase activity was observed in response to a decrease in foliar P concentration of 1.7-1.8 mg P g(DW) (-1).