Chemical modification employed as a means of probing the cell-wall micropore of pine sapwood

The cell-wall micropore network of oven-dried Corsican and Scots pine sapwood has been investigated by reaction with acetic and propionic anhydride in a non-swelling solvent (xylene). No significant reaction occurred, except in the case of reaction between Corsican pine and acetic anhydride. However, when the wood samples were in a swollen state, reaction occurred between the wood and both anhydrides. It is concluded that the cell-wall micropores of Corsican pine are sufficiently large to allow access of acetic anhydride, but not propionic into the unswollen cell wall; however, with Scots pine, both reagents are too large to gain access. A further series of experiments was performed whereby the volumetric changes of pine sapwood samples due to modification with a range of linear chain anhydrides were determined. With Corsican pine, larger volume changes than theoretically predicted were observed, whereas, with the Scots pine samples, the volume changes were close to those predicted. When account is taken of the volume that the reagent occupies within the cell wall, it was found that larger void volumes occurred in the Corsican pine samples. This result indicates that the cell-wall polymeric environment surrounding a reagent molecule is more compliant in the case of Corsican pine.     

Susceptibility of four conifer species to Gremmeniella abietina

G. abietina causes severe dieback in pole-stage stands of Scots pine in Britain. The susceptibility of ll provenances of Scots pine and of three other conifer species was tested by inoculations. Significant differences in susceptibility were found amongst Scots pine provenances, with those originating from a native Scottish pinewood at Loch Maree showing the highest levels of disease. Corsican pine was shown to be the most susceptible species tested although sporulation was greatest on Scots pine. Norway spruce was also shown to be susceptible when inoculated late in the growing season. Infection in lodgepole pine was negligible. High levels of β-phellandrene were tentatively linked with resistance among trees in a single stand of Scots pine.     

Vapor phase reaction of wood with maleic anhydride (I): dimensional stability and durability of treated wood

Reaction between maleic anhydride (MA) and wood specimens was carried out in a vapor phase reaction system. Reaction conditions such as the ratio of supplied MA to wood, initial moisture content, and reaction temperature were optimized. The MA supplied to the reaction system was effectively absorbed by the wood, and a satisfactorily high dimensional stability was achieved even at a low MA/wood ratio. The dimensional stability increased with rising initial moisture content. When the reaction was conducted at an elevated temperature (180°C), high dimensional stability was attained without remarkable weight increase and bulking. The mechanism of dimensional stabilization was discussed on the basis of the dimensional changes at high humidity and during repeated water soaking and drying. It was shown that the dimensional stabilization arises mainly from a decrease of hygroscopicity. When the reaction was conducted at 180°C, the formation of cross-links in the cell wall was apparent. Following the MA treatment, the antifungal property was remarkably enhanced and met the Japanese Industrial Standard K1571. Therefore, MA treatment in the vapor phase is an effective method to attain antifungal properties as well as high dimensional stability with a small amount of nontoxic reagent.     

Structure and abundance of arachnid communities in Scots and lodgepole pine plantations

Between April and September 1994, six plots within Aberfoyle Forest in Scotland, three of Scots (Pinus sylvestris) and three of lodgepole pine (Pinus contorta), were surveyed for spiders and harvestmen, as these are predators of several forest invertebrate pests.A total of 64 species of spider were trapped in Scots and lodgepole pine; 56 in Scots pine and 32 in Lodgepole pine. The most abundant spiders, both in Scots and Lodgepole pine, were Linyphiidae.Scots pine plots were floristically more diverse and structurally more complex than lodgepole pine plots. More species of spiders and harvestmen (midsummer only) were found at ground level in Scots pine than lodgepole pine plots. More species and a greater numbers of spiders were found in the canopy of Scots pine than in that of lodgepole pine. However, no differences in the abundance of harvestmen were found between the canopies of Scots and lodgepole pine. Spider species diversity was greatest in Scots pine.     

Decay threshold of acetylated rattan (Calamus manan) against soft rot

We investigated the resistance of acetylated rattan against soft rot and other soil inhabiting micro-organisms in comparison with wood of beech and Scots pine. Calamus manan of 10 and 13 years old under rubber tree canopy was acetylated to different levels by reaction times (0.25 to 30 hours) and was tested for soft rot decay for 32 weeks. Acetylated rattan at decay protection thresholds of 15.4% and 16.2% weight gain (WG) were fully protected, as shown by both weight loss and strength loss criteria. The static bending properties of untreated rattan decayed by soft rot were significantly lower than for acetylated rattan.     

Relationships between insect diversity and habitat characteristics in plantation forests

A methodological approach to the identification of biodiversity indicators in commercial forest stands is illustrated by analysis of the relationships between syrphid (hoverflies) and carabid (ground beetles) community composition and diversity, and stand structure and field layer vegetation. Data were collected from 12 commercial forest sites encompassing a range of climatic conditions and different crop types (Scots pine, Sitka spruce, Norway spruce and Corsican pine) across the UK. Comparisons were also made between unmanaged semi-natural Scots pine woods and Scots pine plantations. For both syrphids and carabids, no differences in species richness and diversity were recorded between semi-natural stands and plantations; one rare syrphid considered to be restricted to semi-natural pine stands was also found in spruce. Syrphid species diversity and richness was higher in southern spruce sites than in the northern sites. Northern sites had distinctive carabid communities, as did sites in the New Forest, a large ancient woodland in southern England. Of the measured habitat variables, vertical stand structure showed the best correlation with species richness and diversity of both carabids and syrphids. Richness and diversity were less in stands with high vertical cover values for canopy layers. Stands with higher field layer cover supported greater syrphid diversity, but lower carabid diversity. Measures of stand structure could be used as potential indicators of syrphid and carabid diversity, but additional habitat parameters also need to be tested.     

Free amino acid and protein levels,and γ‐glutamyltransferase activity in Pinus sylvestris apical buds and shoots during the growing season

Free amino acid and protein levels, and γ‐glutamyltransferase activity in apical buds and shoots of Scots pine during the growing season. The aim of the study was to obtain the basic information about nitrogen mobilization needed in carrying out studies on the optimal nitrate and ammonium ratios in the metabolism of Scots pine (Pinus sylvestris L.). Considerable seasonal changes in the concentrations of free amino acids and other ninhydrin‐positive low molecular‐weight compounds were observed in the buds and shoots of Scots pine. 43 different amino compounds were identified, the concentrations of arginine, glutamine, glutamic acid, γ‐aminobutyric acid, alanine and aspartic acid being highest at the break of dormancy. The amounts of certain amino compounds decreased during the growing season, those of arginine, ethanolamine and various ammonium compounds in particular. The amount of glutamic and aspartic acids, glycine, alanine and γ‐aminobutyric acid, however, remained relatively constant. The protein concentration and the specific activity of γ‐glutamyltransferase increased towards the end of the growing period. Slight differences were found between the fertilized trees and the control trees.     

Glucanohydrolase enzyme activity in embryos of Scots,Corsican and lodgepole pines infected in vitro with Heterobasidion annosum

Embryos of Scots pine (Pinus sylvestris), Corsican pine (P. nigra var. maritima) and Lodgepole pine (P. contorta) were infected in vitro with Heterobasidion annosum and the activities of 1,3-β-glucanase, chitinase and β-glucosidase were monitored. Both the infected and control embryos of the three pine species showed the presence of the enzymes. Statistical analysis showed no significant increase (P > 0.05) in the levels of 1,3-β-glucanase in infected embryos for all the three pine species. However, there was a significant increase in levels of chitinase in infected embryos of P. contorta and P. nigra var. maritima and also in levels of β-glucosidase in infected embryos of all three of the pine species. The study has therefore revealed that constitutive levels of glucanohydrolase enzymes exist in the embryos of the three pine species and that the induction of chitinase and β-glucosidase was a factor in the response of the embryos to stimulation by H. annosum infection.     

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.     

Impact of decreasing throughfall depositions on soil solution chemistry at coniferous monitoring sites in northern Belgium

Impact of decreasing throughfall depositions on soil solution chemistry at three depths was examined in a Corsican pine (Pinus nigra Arn. ssp. laricio Poiret) stand in Ravels and a Scots pine (Pinus sylvestris L.) stand in Brasschaat within a time-span of 6 years (1992–1997). At Ravels depositions of sulphate, ammonium, nitrate, calcium, magnesium and sodium decreased by 463, 468, 169, 121, 63 and 169 mol_c ha⁻¹ per annum. At Brasschaat deposition reduction rates of sulphate, nitrate, calcium and magnesium were 198, 127, 134 and 46 mol_c ha⁻¹ per annum. In both stands the substantial reductions in inorganic nitrogen deposition were followed by a decline of nitrate fluxes leaving the forest floor and with the seepage water. The decrease of sulphate deposition since 1992 was only manifested by a decline in sulphate concentrations and fluxes during the first half of the measuring period. The increase at the end of the period could be ascribed to the fact that sulphate adsorption which was important between 1993 and 1995 finally ceased. A significant drop of calcium concentrations was discernible at both plots. Magnesium and potassium levels did only fall significantly at Brasschaat. The concentrations of base cations were largely driven by the concentrations of their counter anions, but were influenced by the base cation throughfall fluxes as well. Although a substantial reduction of throughfall acidity occurred, no improvement of the soil water acidity (acid neutralising capacity (ANC)) was noticed. ANC and molar Ca/Al followed decreasing trends at all depths in both plots. For ANC the decline was significant for the topsoil in Ravels. The decrease of molar cation Ca/Al ratio was significant for two out of three depths at both sites. The decrease was due to the drop in calcium concentrations and the raise of Al_tot in some horizons.     

Kinetics of wood-anhydride reactions: a novel approach

The reaction kinetics of wood (Corsican pine) with acetic, propionic, succinic, and hexenyl succinic anhydride dissolved in pyridine were investigated. To determine reaction kinetics, anhydride depletion of the reactive liquid was monitored. This approach allows to collect a complete sequence of kinetic data from a single experiment. The results carry information on reaction kinetics without allowing for a precise determination of reaction turnover vs time. For all anhydrides investigated, the reaction kinetics of wood samples of different size (300 μm–4 mm) do not depend on sample size, hence the reaction is activation- not diffusion-controlled. All results are consistent with a remarkably simple model of purely activation-controlled reaction kinetics with a single activation energy. Activation energies were determined. The result for propionic anhydride agrees with an earlier investigation based on weight gain of reacted wood. Received 16 March 1999     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Dynamics of some mineral nutrients and heavy metals in decomposing forest litter

The dynamics of three major mineral nutrients (K, Ca, Mg) and six heavy metals (Fe, Mn, Zn, Cu, Pb and Cd) were studied in decomposing natural, unpolluted litter in two forest types: Scots pine needle litter in a pure Scots pine forest and oak‐hornbeam leaf litter in a mixed oak‐hornbeam stand. Of the mineral nutrients, only K in the oak‐hornbeam system showed a significant decrease in concentration during the course of litter decomposition. For Ca and Mg in both systems and for K in the Scots pine system no clear relation between concentration and accumulated litter‐mass loss was observed. On the other hand, for Fe, Zn, Pb and Cd at both sites, as well as for Mn in the oak‐hornbeam stand and Cu in the Scots pine stand highly significant positive relations were found between heavy metal concentrations and accumulated mass loss. A decrease in concentration was noted only in the case of Mn in the Scots pine system. No clear pattern was found for Cu in the oak‐hornbeam stand. Concentrations of heavy metals in the decomposing litter reached levels at or above those reported to retard decomposition in other studies. At the oak‐hornbeam stand increased also the absolute amounts of Fe, Zn, Pb and Cd.     

A new multivariate approach to foliar analyses for diagnostic purposes in mature Scots pine stands

The relationship between the needle dry weight and the contents of particular nutrients in pine needles is studied using multiple regression and path coefficient analysis. By joint use of all the information thus obtained it is possible to reach an objective assessment of the nutrient status of mature pine stands and to plan adequate fertilization without recourse to uncertain critical nutrient levels and to the difficult-to-interpret tree growth response. An example is presented of the application of this approach to a 72-year-old Scots pine stand.     

Infection experiments with Gremmeniella abietina on seedlings of Norway spruce and Scots pine

Conidia of Gremmeniella abietina infected and caused disease symptoms in annual shoots of both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings. In Norway spruce shoots the infection remained largely latent, with only a few seedlings showing symptoms. Mycelial growth inside the shoots was faster in Scots pine than in Norway spruce and was favoured by low temperature in both hosts. The shoots of Norway spruce seedlings had higher endophyte populations than those of Scots pine, and the populations were decreased by low temperatures. Reductions in the normal epiphytic or endophytic flora by acid mist treatments seemed to favour the development of G. abietina.     

Mixed acylation of Scots pine sawdust and impact on hydrophobicity

Abstract

Acetic–fatty esters of Scots pine sawdust (SPS) were obtained by reaction of SPS with mixtures containing acetic–fatty anhydrides and no solvent or catalyst. Such mixtures were synthesized by reaction between a carboxylic acid and acetic anhydride. The global reaction of acetic anhydride and a fatty acid yields at equilibrium a mixture of five compounds: acetic–fatty anhydride, acetic anhydride, fatty acid, acetic acid and fatty anhydride. The influence of temperature, molar ratio, reaction time and length of the fatty chain on the esterification and on the ratio of grafted acetyl/fatty acyl was investigated. Static contact angles (CAs) with water were measured over 5 min. CA values were dependent on the fatty acyl content and independent of the acetyl content. The minimum ester content of the oleoyl group required to obtain permanent water repellency was 25 mmol kg^?1. Water vapour adsorption measurements indicated that in contrast to water repellency, hydrophobicity to water vapour was correlated with the total mass acyl content.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Genetic variation and divergence in Scots pine (Pinus sylvestris L.) within its natural range in Italy

Twenty-one populations of Scots pine sampled over the entire Italian range of the species were analysed for genetic variation scored at nine nuclear SSR markers. The main aim of the work was to find genetic features useful for conservation management, namely allelic composition, gene diversity and differentiation. High levels of intra-population variability were scored. The only population sampled in the Apennines gave the lowest values, confirming the genetic erosion undergone in the Scots pine remnants in this area. A low level of genetic variability was also scored for populations from the Po valley and hills of Piedmont. Most genetic diversity was found within populations, while only a small amount occurred among them (F _ST?=?0.058). Both Bayesian clustering and sPCA analysis showed a East–West subdivision, notwithstanding the unclear position of populations from the Po valley. The population from the Apennines was always clearly separated from the others. The results are discussed in terms of post-glacial recolonisation, as well as for defining genetically homogeneous regions for Scots pine in Italy. The management of genetic resources could benefit from the identification of such ‘gene zones’, thereby avoiding the use of non-local reproductive material for plantations, which can represent one of the most important reasons for failure of reforestation. In addition, the assessment of the biogeographic genetic structure by neutral markers is a prerequisite for disentangling the influence of selectively neutral and non-neutral processes on the distribution of adaptive genetic variability.     

Acetylation of wood in combination with polysiloxanes to improve water-related and mechanical properties of wood

Scots pine sapwood was acetylated with ethyltriacetoxysilane using acetic acid as a solvent and sulfuric acid as a catalyst. A weight percent gain (WPG) of 14 % and cell wall bulking of 7 % were obtained after 5 h of reaction time. Pine specimens were acetylated with acetic anhydride in the presence of 1 % ethyltriacetoxysilane, dihydroxy-functional siloxane, acetoxy-functional siloxane, amino-functional siloxane and non-functional siloxane, respectively. Acetoxy-functional siloxane induced the greatest reduction in water uptake with a water repellent effectiveness after 24 h of up to 62 % as compared to acetylated wood. WPG and cell wall bulking increased compared to solely acetylated wood with increasing concentrations of acetoxy-functional siloxane in acetic anhydride; anti-shrink efficiency, however, did not increase. Fungal resistance of pine sapwood and beech as well as mechanical strength properties did not change when 20 % acetoxy-functional siloxane was added to acetic anhydride compared to solely acetylated specimens.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.