Colour of larch heartwood and relationships to extractives and brown-rot decay resistance

Larch heartwood is appreciated for its good mechanical properties, its colour and its texture, and it is often used outdoors because of its natural durability (decay resistance). In this study the colour of larch heartwood was studied in relation to extractives and decay resistance, with the aim to estimate durability of larch heartwood from its colour. On a total of 293 trees colour in the CIE L*a*b* space (L* lightness, a* red/green axis, b* yellow/blue axis), extractives content (acetone and hot-water extractives, amount of phenolics) and the brown-rot decay resistance were determined. For calculating the relative decay resistance ( x), mass loss after inoculation for 16 weeks with two fungi [ Coniophora puteana (Schum.ex.Fr.) Karst., Poria placenta (Fr.) Cke, European standard EN 113] of larch heartwood samples was compared to Scots pine ( Pinus sylvestris L) sapwood reference samples (EN 350-1). Different species [Japanese larch ( Larix kaempferi Lamb.), Hybrid larch (Larix deciduax L. kaempferi) and European larch ( L. decidua Mill.)], provenances and age classes (38-year, >150-year) were included. Japanese larch heartwood turned out to be significantly more reddish (higher a*-values) compared to the European larch provenances. Reddishness of the hybrids was intermediate. The red hue (+a*) was strongly correlated with the amount of phenols ( r =0.84) and decay resistance ( r =0.63) and therefore suitable for prediction of both parameters. The results suggest that colour measurements of larch heartwood could be of benefit in tree breeding programs and for an optimised utilization of larch timber.     

Investigations on natural durability of important European wood species against wood decay fungi. Part 1: Laboratory tests

The durability of heartwood from European larch, Sessile oak and Scots pine was tested in laboratory against wood decay basidiomycetes and soft rot. The durability test was performed according to CEN/TS 15083-1:2005 with Coniophora puteana, Oligoporus (Poria) placenta and Trametes (Coriolus) versicolor. CEN/TS 15083-2:2005 was applied in order to test the timber with a bioactive soil substrate against soil rotting organisms. For Sessile oak, a very high durability against basidiomycetes (DC 1) was found in contrast to a low durability (DC 4) against soft rot fungi. Furthermore the results indicated that the durability of European larch and Scots pine is slightly deteriorated after leaching according to EN 84:1997. Beside this a minor influence of raw density on mass loss was detected for larch.     

Resistance of Scots pine wood to Brown-rot fungi after long-term forest fertilization

The susceptibility of Scots pine (Pinus sylvestris L.) sap- and heartwood against the wood decaying brown-rot fungus (Coniophora puteana) was investigated after long-term forest fertilization at three different sites in central Finland. Different wood properties: wood extractives, wood chemistry, and wood anatomy were used to explain sap- and heartwood decay. Scots pine sapwood was more susceptible to decay than its heartwood. In one site, sapwood seemed to be more resistant to wood decay after forest fertilization whereas the susceptibility of heartwood increased. Significant changes in the sapwood chemistry were found between treatment and sites, however, no relationship between wood chemistry and wood decay was observed in the factor analysis. The results of this study show that there was an inconsistent relationship between decay susceptibility and fertilization and the measured physical and chemical attributes of the wood were not consistently correlated with the decay rate.     

Durability of five native Argentine wood species of the genera <Emphasis Type="Italic">Prosopis</Emphasis> and <Emphasis Type="Italic">Acacia</Emphasis> decayed by rot fungi and its relationship with extractive content

The natural durability of four Argentinean species of Prosopis and one of Acacia was evaluated in laboratory tests, according to European standards, using three brown rot and one white rot fungi. These tests were complemented by assessing the wood chemical composition. All the species were from moderately slightly durable to very durable (classes 4–1), and in all cases the heartwood was the most resistant to fungal attack. Chemical extractives content (organic, aqueous, tannic and phenolic) was higher in the heartwood. However, species durability was not related to extractive contents nor with wood density. Instead, it is possible that extractives could contribute to natural durability in different ways, including the effects related to the antioxidant properties of some of them.     

Chemical factors affecting the brown-rot decay resistance of Scots pine heartwood

The cell wall chemistry (amount of hemicellulose, f-cellulose, and total lignin) and the concentration of extractives (total acetone-soluble extractives, resin acids, pinosylvins and the total phenolics quantified as tannin acid equivalents) were studied in brown-rot resistant and susceptible juvenile heartwood of Scots pine (Pinus sylvestris L.). The study material consisted of a total of 18 trees from two 34-year-old progeny trials at Korpilahti and Kerimäki. The trees were selected from among 783 trees whose decay rate had previously been screened in a laboratory test using a brown-rot fungus, Coniophora puteana. Samples from neither location showed any significant difference in the concentration (mg/cm³) of hemicellulose, f-cellulose and total lignin between the decay resistant and susceptible trees. At both locations only the concentration of total phenolics was higher in the decay-resistant heartwood than in the decay-susceptible heartwood. At Korpilahti, the amount of acetone-soluble extractives and the concentration of pinosylvin and its derivatives were higher in the resistant than in the susceptible trees.     

Heartwood extractives and lignin content of different larch species (<Emphasis Type="Italic">Larix </Emphasis>sp.) and relationships to brown-rot decay-resistance

The extractive content of lignin and the brown-rot decay-resistance against Coniophora puteana and Poria placenta were studied in larch heartwood from different species and origin (Larix decidua var. decidua, L. decidua var. sudetica, L. kaempferi , L. × eurolepis). The study material consisted of 106 trees from a 39-year old provenance trial in France. The hot-water-soluble extractives were very variable (from5.66% to 20.50% of dry weight), but there was no significant variation between the investigated species and origins. In contrast, acetone extractives, the total amount of phenolics and lignin showed significant differences. The concentration of phenolics and lignin was significantly higher in L. kaempferi and in L. × eurolepis than in L. decidua. The total phenolics content was strongly correlated with decay-resistance in all investigated larch origins. A higher concentration of phenolics goes hand in hand with higher decay resistance and phenolics might therefore be a promising parameter to rapidly evaluate the level of decay-resistance in larch.     

Performance of the potentially invasive Siberian moth Dendrolimus superans sibiricus on coniferous species in Europe

1 The native range of the Siberian moth extends from the Pacific Ocean (Russian Far East, Japan and Northern Korea) across Siberia, Northern China and Mongolia to the Ural Mountains. At the beginning of the 21st Century, this species was documented west of the Ural Mountains in the Republic of Mari El, indicating range extension toward the west.
2 The Siberian moth has recently been suggested for regulation as a quarantine pest for European and Mediterranean Plant Protection Organization member countries. However, no specific report on European host plants for this pest has been published so far.
3 In the present study, larval host plant choice and performance was tested for the first time on coniferous tree species that are widely distributed and of commercial value in Europe.
4 Based on dual-choice tests on neonates and mortality, developmental duration and relative growth rates of the first- to sixth-instar larvae, we found European larch Larix decidua to be the most suitable host for the moth larvae, whereas European black pine Pinus nigra and Scots pine Pinus sylvestris were the poorest hosts. The remaining conifer species tested, European silver fir Abies alba , Nordmann fir Abies nordmanniana , and Norway spruce Picea abies , were intermediate host plants. Douglas-fir Pseudotsuga menziesii , originating from North America, was chosen by the larvae to the same extend as European larch, and was also highly suitable for larval development.
5 If the moth is introduced to European countries, it will become damaging in stands of European larch and Douglas-fir, mixed stands of fir and spruce; however, it will be less damaging in forests dominated by two-needle pines.
6 We predict that Dendrolimus superans sibiricus will be able to survive and develop on the main European coniferous tree species, including non-native coniferous tree species, resulting in severe damage to large areas of forests.     

Differences in nitrate and ammonium uptake between Scots pine and European larch

In forest soils, ammonium is usually the predominant form of inorganic nitrogen. However, the capacity of trees to utilize both NO₃ ^- and NH₃ ⁺ may provide greater flexibility in responding to changes of nitrogen supply from the environment. Such capacity has been studied in seedlings of Scots pine (Pinus sylvestris L.) and European larch (Larix decidua Mill.) grown in the presence or absence of either nitrate or ammonium. Nitrate-induced plants showed a higher nitrate uptake rate than non-induced plants; this difference was almost negligible after 24 h of exposure to NO₃ ^-. Ammonium uptake in both species was consistently higher than that of nitrate, regardless of prior nitrogen provision. In both nutrient conditions, larch showed a more efficient transport system in comparison with Scots pine, with higher ammonium and nitrate uptake rates in both induced and non-induced plants. This was consistent also with the activity of nitrate reductase, measured in vivo in roots and leaves. This revised version was published online in June 2006 with corrections to the Cover Date.     

Estimating missing sapwood rings in three European gymnosperm species by the heartwood age rule

Precise dating of the year of felling is one intended outcome of dendrochronology. However, occasionally some or all sapwood rings might be missing, either due to deterioration or because they were carved off, or for some other reason. Consequently, while heartwood is preserved, sapwood might be fully or partially missing. In such cases, the year of felling must be estimated by adding a suitable number of sapwood rings. A heartwood age rule (HAR) has been advocated for Scots pine and adapted to European larch and Cembra pine, implying a linear relationship between sapwood ring count and the square root of heartwood ring count, largely irrespective of position in the stem. The same rule applied to all observations of a species, irrespective of silviculture, location or fertility of the growth site. Scots pine had twice or thrice as many sapwood rings as Cembra pine, which had 10% more rings than larch. The magnitude of model residuals was proportional to estimated sapwood ring count. Relative residuals were roughly normally distributed. To be applicable in Bayesian modelling in dendrochronology analyses, detailed information on model errors has been provided.     

The effect of water soluble Scots pine (Pinus sylvestris L.) and Sitka spruce [Picea sitchensis (Bong.) carr.] heartwood and sapwood extracts on the growth of selected Trichoderma species

Extractable materials from some timber species have been identified which prevent wood decay; however, little has been reported on the effect(s) of such materials against mould species that colonize timber. With increasing interest in the use of Trichoderma species, both as agents of permeability enhancement and biological control, more information is required on how chemical components within fresh and processed timber influence growth of Trichoderma. Fresh and dried samples of Scots pine and Sitka spruce sapwood and heartwood were leached in a Soxhlet apparatus and the resulting extract was combined with malt extract agar and inoculated with Trichoderma. Trichoderma isolates were inhibited to varying degrees by extractives removed from fresh and dried heartwood of the two timbers. Growth on sapwood extractives, however, showed a lesser degree of inhibition. The implications of the results for applications of Trichoderma in timber are discussed.     

Endurance of larch forest ecosystems in eastern Siberia under warming trends

The larch (Larix spp.) forest in eastern Siberia is the world's largest coniferous forest. Its persistence is considered to depend on near‐surface permafrost, and thus, forecast warming over the 21st century and consequent degradation of near‐surface permafrost is expected to affect the larch forest in Siberia. However, predictions of these effects vary greatly, and many uncertainties remain about land – atmosphere interactions within the ecosystem. We developed an integrated land surface model to analyze how the Siberian larch forest will react to current warming trends. This model analyzed interactions between vegetation dynamics and thermo‐hydrology, although it does not consider many processes those are considered to affect productivity response to a changing climate (e.g., nitrogen limitation, waterlogged soil, heat stress, and change in species composition). The model showed that, under climatic conditions predicted under gradual and rapid warming, the annual net primary production of larch increased about 2 and 3 times, respectively, by the end of the 21st century compared with that in the previous century. Soil water content during the larch‐growing season showed no obvious trend, even when surface permafrost was allowed to decay and result in subsurface runoff. A sensitivity test showed that the forecast temperature and precipitation trends extended larch leafing days and reduced water shortages during the growing season, thereby increasing productivity. The integrated model also satisfactorily reconstructed latitudinal gradients in permafrost presence, soil moisture, tree leaf area index, and biomass over the entire larch‐dominated area in eastern Siberia. Projected changes to ecosystem hydrology and larch productivity at this geographical scale were consistent with those from site‐level simulation. This study reduces the uncertainty surrounding the impact of current climate trends on this globally important carbon reservoir, and it demonstrates the need to consider complex ecological processes to make accurate predictions.     

Naturally durable heartwood: evidence for a proposed dual defensive function of the extractives

We previously proposed that extractives in highly durable heartwood may protect wood against fungal colonization and subsequent degradation by dual mechanisms: the extractives have some fungicidal activity and are also free radical scavengers (antioxidants). In short-term laboratory decay tests using two different wood species and decay fungi, the antioxidant 2,6-dimethyl-di-tert-butyl-4-methylphenol (BHT) alone had little or no preservative effect. In contrast, the combination of BHT with different organic commercial biocides always showed an increase in efficacy compared to the organic biocide alone. Consequently, we conclude that the combination of a commercial antioxidant and biocide is synergistic. This implies that extractives may protect wood by more than simply being fungicidal.     

The effect of single tree species on soil microbial activities related to C and N cycling in the Siberian artificial afforestation experiment

The effects of grassland conversion to forest vegetation and of individual tree species on microbial activity in Siberia are largely unstudied. Here, we examined the effects of the six most commonly dominant tree species in Siberian forests (Scots pine, spruce, Arolla pine, larch, aspen and birch) on soil C and N mineralization, N₂O-reduction and N₂O production during denitrification 30 years after planting. We also documented the effect of grassland conversion to different tree species on microbial activities at different soil depths and their relationships to soil chemical properties. The effects of tree species and grassland conversion were more pronounced on N than on C transformations. Tree species and grassland conversion did significantly alter substrate-induced respiration (SIR) and basal respiration, but the differences were not as large as those observed for N transformations. Variances in SIR and basal respiration within species were markedly lower than those in N transformations. Net N mineralization, net nitrification, and denitrification potential were highest under Arolla pine and larch, intermediate under deciduous aspen and birch, and lowest beneath spruce and Scots pine. Tree species caused similar effects on denitrification potential, net N mineralization, and net nitrification, but effects on N₂O reduction rate were idiosyncratic, indicating a decoupling of N₂O production and reduction. We predict that deciduous species should produce more N₂O in the field than conifers, and that Siberian forests will produce more N₂O if global climate change alters tree species composition. Basal respiration and SIR showed inverse responses to tree species: when basal respiration increased in response to a given tree species, SIR declined. SIR may have been controlled by NH₄ ⁺ availability and related therefore to N mineralization, which was negatively affected by grassland conversion. Basal respiration appeared to be less limited by NH₄ ⁺ and controlled mostly by readily available organic C (DOC), which was higher in concentration under forests than in grassland and therefore basal respiration was higher in forested soils. We conclude that in the Siberian artificial afforestation experiment, soil C mineralization was not limited by N.     

Phylogeography of Scots pine in Europe and Asia based on mtDNA polymorphisms

We analyzed mitochondrial DNA polymorphisms to search for evidence of the genetic structure and patterns of admixture in 124 populations (N = 1407 trees) across the distribution of Scots pine in Europe and Asia. The markers revealed only a weak population structure in Central and Eastern Europe and suggested postglacial expansion to middle and northern latitudes from multiple sources. Major mitotype variants include the remnants of Scots pine at the north-western extreme of the distribution in the Scottish Highlands; two main variants (western and central European) that contributed to the contemporary populations in Norway and Sweden; the central-eastern European variant present in the Balkan region, Finland, and Russian Karelia; and a separate one common to most eastern European parts of Russia and western Siberia. We also observe signatures of a distinct refugium located in the northern parts of the Black Sea basin that contributed to the patterns of genetic variation observed in several populations in the Balkans, Ukraine, and western Russia. Some common haplotypes of putative ancient origin were shared among distant populations from Europe and Asia, including the most southern refugial stands that did not participate in postglacial recolonization of northern latitudes. The study indicates different genetic lineages of the species in Europe and provides a set of genetic markers for its finer-scale population history and divergence inference.     

Seasonal changes of pinosylvin distribution in the sapwood/heartwood boundary of Pinus sylvestris

Changes in pinosylvin concentration and distribution across the sapwood/heartwood boundary were studied on Scots pine (Pinus sylvestris L.) tree stems to detect seasonal activity in heartwood formation. Pinosylvin concentrations were measured with FT-(NIR) Raman spectroscopy for a total of 96 trees equally distributed on 16 different sampling occasions. In another experiment, cores were sampled every month from six individual Scots pine trees from June to October and analysed for pinosylvin. There was a great between-tree variation in concentration and spatial distribution of pinosylvin. No seasonal trend in the distribution pattern or concentration of pinosylvin was found. Therefore, the results indicate that there is no specific period during the year when heartwood is formed. Received: 12 February 1998 / Accepted: 3 May 1999     

Wood durability of home-garden teak against brown-rot and white-rot fungi

Natural decay resistance of teak wood grown in home-garden forestry and the factors influencing decay resistance were determined in comparison with that of a typical forest plantation. Accelerated laboratory tests were conducted on 1800 wood samples drawn from 15 trees of three planted sites. Analysis of variance based on a univariate mixed model showed that planted site, fungal species, and their interaction terms were important sources of variation in decay resistance. With increasing decay resistance from centre to periphery of the heartwood, radial position was a critical factor and the interaction effect of fungal species × radial position was significant in influencing the durability. No significant differences were found in decay resistance either between the opposite radii or due to the various possible interaction terms of radii with the site, fungal species and radial position. There were significant differences in decay resistance against brown-rot fungi between wet and dry sites of home-garden teak although differences against white-rot fungi were non-significant among the three planted sites. Polyporus palustris was the more aggressive brown-rot fungus than Gloeophyllum trabeum. The higher susceptibility of wet site home-garden teak to brown-rot decay was associated with a paler colour of the wood and lower extractive content.     

Tall shrub and tree expansion in Siberian tundra ecotones since the 1960s

Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest‐tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in 11, widely distributed Siberian ecotonal landscapes by comparing very high‐resolution photography from the Cold War‐era ‘Gambit’ and ‘Corona’ satellite surveillance systems (1965–1969) with modern imagery. We also analyzed within‐landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of 11 ecotones. In northwest Siberia, alder (Alnus) shrubland cover increased 5.3–25.9% in five ecotones. In Taymyr and Yakutia, larch (Larix) cover increased 3.0–6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice‐rich permafrost. In Chukotka, the total cover of alder and dwarf pine (Pinus) increased 6.1% within one ecotone and was little changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned‐ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid‐1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape scale. Our results indicate that extensive changes can occur within decades in moist, shrub‐dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch‐dominated ecotones of central and eastern Siberia.     

Genetic improvement of the chemical composition of Scots pine (Pinus sylvestris L.) juvenile wood for bioenergy production

Chemical composition is one of the key characteristics that determines wood quality and in turn its suitability for different end products and applications. The inclusion of chemical compositional traits in forest tree improvement requires high‐throughput techniques capable of rapid, non‐destructive and cost‐efficient assessment of large‐scale breeding experiments. We tested whether Fourier‐transform infrared (FTIR) spectroscopy, coupled with partial least squares regression, could serve as an alternative to traditional wet chemistry protocols for the determination of the chemical composition of juvenile wood in Scots pine for tree improvement purposes. FTIR spectra were acquired for 1,245 trees selected in two Scots pine (Pinus sylvestris L.) full‐sib progeny tests located in northern Sweden. Predictive models were developed using 70 reference samples with known chemical composition (the proportion of lignin, carbohydrates [cellulose, hemicelluloses and their structural monosaccharides glucose, mannose, xylose, galactose, and arabinose] and extractives). Individual‐tree narrow‐sense heritabilities and additive genetic correlations were estimated for all chemical traits as well as for growth (height and stem diameter) and wood quality traits (density and stiffness). Genetic control of the chemical traits was mostly moderate. Of the major chemical components, highest heritabilities were observed for hemicelluloses (0.43–0.47), intermediate for lignin and extractives (0.30–0.39), and lowest for cellulose (0.20–0.25). Additive genetic correlations among chemical traits were, except for extractives, positive while those between chemical and wood quality traits were negative. In both groups (chemical and wood quality traits), correlations with extractives exhibited opposite signs. Correlations of chemical traits with growth traits were near zero. The best strategy for genetic improvement of Scots pine juvenile wood for bioenergy production is to decrease and stabilize the content of extractives among trees and then focus on increasing the cellulose:lignin ratio.     

Plasticity in gas‐exchange physiology of mature Scots pine and European larch drive short‐ and long‐term adjustments to changes in water availability

Adjustment mechanisms of trees to changes in soil‐water availability over long periods are poorly understood, but crucial to improve estimates of forest development in a changing climate. We compared mature trees of Scots pine (Pinus sylvestris) and European larch (Larix decidua) growing along water‐permeable channels (irrigated) and under natural conditions (control) at three sites in inner‐Alpine dry valleys. At two sites, the irrigation had been stopped in the 1980s. We combined measurements of basal area increment (BAI), tree height and gas‐exchange physiology (Δ¹³C) for the period 1970–2009. At one site, the Δ¹³C of irrigated pine trees was higher than that of the control in all years, while at the other sites, it differed in pine and larch only in years with dry climatic conditions. During the first decade after the sudden change in water availability, the BAI and Δ¹³C of originally irrigated pine and larch trees decreased instantly, but subsequently reached higher levels than those of the control by 2009 (15 years afterwards). We found a high plasticity in the gas‐exchange physiology of pine and larch and site‐specific responses to changes in water availability. Our study highlights the ability of trees to adjust to new conditions, thus showing high resilience.     

Fixation,leachability, and decay resistance of wood treated with some commercial extracts and wood preservative salts

Beech (Fagus orientalis) and Scots pine (Pinus sylvestris) wood blocks were treated with some commercial extracts as well as water-based wood preservative salts at various concentrations to increase retention and fixation. The penetration, fixation, and antifungal properties of different treatment solutions were compared with statistical analysis. Retention levels of solutions were generally higher for Scots pine wood than beech wood. The highest retention levels were seen in wood treated with sumac leaf extract and oak valonia extract. Leaching tests indicated that both wood types treated with sumac extracts showed higher retention levels than wood treated with the other fruit and bark extract solutions. Adding 1% water-based wood preservative salts to valonia and sumac leaf extracts increased the retention levels. Concentrations of more than 1% did not contribute to retention either individually or with salt additions. Three percent and higher concentrations of wood-preserving salts accelerated and increased the amount of leaching. The results showed that the extract alone was resistant to leaching. Mycological tests showed that all extractives were significantly effective against wood decay.