Thioacidolytic analysis of residual lignins isolated from the chemical pulps of spruce (Picea abies) and beech (Fagus sylvatica) wood

Eight different residual lignins isolated from unbleached chemical pulps [sulfite, kraft, alkaline sulfite-anthraquinone-methanol (ASAM), soda/AQ/MeOH] of spruce and beech were characterized using gel permeation chromatography (GPC), thioacidolysis, and desulfurization to determine average molecular weight, amounts of uncondensed β-O-4 linkages, and dimeric linkage patterns, respectively. The total amounts of G-CHSEt-CHSEt-CH₂SEt and S-CHSEt-CHSEt-CH₂SEt were markedly reduced in residual lignins to 40% to 80% of the values for the corresponding milled wood lignins (MWLs). The number of dimeric units determined by thioacidolysis and desulfurization of the residual lignins was decreased by onehalf to one-fifth compared to the MWLs. Among the diverse types of dimeric units, reduction of β-1 and β-5 units was significant in most of the residual lignins, with the exception of beech ASAM residual lignin. Compared to beech MWL, 40% more C6–C3 main monomers were detected, whereas the relative composition of the dimeric units in the beech residual lignin was very similar to that in the beech MWL. The average molecular weights of residual lignins were less than those of the MWLs. However, the average molecular weights of the spruce kraft and soda residual lignins were determined to be higher than those of the corresponding MWLs.     

Improving estimations of maximal stand density by combining Reineke’s size-density rule and the yield level, using the example of spruce (Picea abies (L.) Karst.) and European Beech (Fagus sylvatica L.)

? The Reineke Stand density rule relating stem numbers to the quadratic mean diameter is generally used as a reference for modelling maximal stand density.

? The linearity of this relationship after double logarithmic transformation is generally assumed, but it must be questioned for untouched stands and stands with a conventional thinning regime. Curvilinearity is demonstrated for some spruce and beech stands in Switzerland and shown to be statistically representative. This relationship is independent of the site index. It can be interpreted as a change in mortality in young stages mainly due to competition and in older stages more to ageing.

? A more accurate estimation of the maximal stand density needs to take into account the important variation around the mean course, known as the yield level. A simple method to assess the yield level of any stand regardless of whether it is thinned or not is presented, based on estimating the effect of a stand opening on the basal area.

     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

Spatial and temporal variability of natural regeneration in a temperate old-growth forest

? The natural regeneration of shade-tolerant tree species is characterized by large spatial heterogeneity and temporal variability. The mechanisms producing those patterns are still poorly understood and the knowledge of long-term fluctuations in regeneration processes is very limited.

? We used data from long-term study plots in an old-growth stand dominated by European beech and silver fir to address three questions: (1) Is a tendency towards clumping in seedlings associated with a particular spatial scale? (2) Are the spatial patterns in seedlings constant over time? (3) Is the distribution of seedlings related to light intensity?

? Over the study period the seedling density varied from 2.5 to 6 ind./m². Stronger fluctuations occurred at a fine scale. Seedlings were clumped at distances ranging from a few centimeters to a few meters. The distribution of places with high seedling densities was relatively constant over time, especially in beech. Positive but weak rank correlations were found between light intensity and densities of beech and fir seedlings.

? Despite large fluctuations in seedlings densities, caused by mast-seeding, a permanent seedling bank was maintained. Very pronounced clumping of tree seedlings found in this study was only partially explained by differences in light intensity.

     

Temporal trends in the foliar nutritional status of the French, Walloon and Luxembourg broad-leaved plots of forest monitoring

? Two decades after the launching of the monitoring program of forest ecosystems in Europe (ICP forests), a unique data set is now available regarding the foliar nutritional status of the main broad-leaved species growing in much-diversified sites (soil, climate).

? This study focuses on the foliar concentration time series (1993 to 2005) of the French (RENECOFOR), Walloon and Luxemburg broad-leaved plots. The aim is to show long-term trends while taking the inter-plot and inter-annual variability into account.

? Two kinds of statistical processing were used to analyze the data on foliar chemistry: principal component analysis (PCAs) and linear mixed models. In general, the main temporal trends reveal a decrease in the foliar P concentration and a decrease in the foliar Ca concentration, except for European beech on acid soils.

? These temporal patterns confirm the trends previously observed and could be explained by the joint actions of several processes that influence tree nutrition in the long term: tree age, N and S deposition, harvesting and climate.

     

Comparison of model forms for estimating stem taper and volume in the primary conifer species of the North American Acadian Region

? The performance of ten commonly used taper equations for predicting both stem form and volume in balsam fir [Abies balsamea (L.) Mill], red spruce[Picea rubens (Sarg.)], and white pine[Pinus strobus (L.)] in the Acadian Region of North America was investigated.

? Results show that the Kozak (2004) and Bi (2000) equations were superior to the other equations in predicting diameter inside bark for red spruce and white pine, while the Valentine and Gregoire (2001) equation performed slightly better for balsam fir.

? For stem volume, the Clark et al. (1991) equation provided the best predictions across all species when upper stem diameter measurements were available, while the Kozak (2004) and compatible taper equation of Fang et al. (2000) performed well when those measurements were unavailable.

? The incorporation of crown variables substantially improved stem volume predictions (mean absolute bias reduction of 7–15%; root mean square error reduction of 10–15%) for all three species, but had little impact on stem form predictions.

? The best taper equation reduced the predicted root mean square error by 16, 39, and 45% compared to estimates from the widely used Honer (1965) regional stem volume equations for balsam fir, red spruce, and white pine, respectively.

? When multiple taper equations exist for a certain species, the use of the geometric mean of all predictions is an attractive alternative to selecting the “best” equation.

     

The importance of canopy-derived dissolved and particulate organic matter (DOM and POM) — comparing throughfall solution from broadleaved and coniferous forests

? Dissolved organic matter (DOM) and its main constituents carbon (DOC) and nitrogen (DON) represent an important part of the C and N cycles in forest ecosystems. Although many investigations have been addressing this issue, the knowledge on particulate organic matter (0.45 μm < POM < 500 μm) dynamics, its origin and involvement in organic matter cycling in forest ecosystems is still imperfect.

? In this paper, we report on dissolved and particulate organic carbon and nitrogen fractions in throughfall solutions collected from a broadleaved and coniferous forest stand in Central Germany. Over a period of 2.5 y (2005–2007) we followed the concentrations and fluxes of DOM and POM at a mature beech (Fagus sylvatica L.) and a Norway spruce (Picea abies L.) forest site. Bulk and throughfall precipitation were sampled in weekly (2005) and fortnightly (2006–2007) intervals and analyzed for dissolved (< 0.45 μm, filtered) and total (< 500 μm, unfiltered) amounts of organic carbon (DOC, TOC, POC) and nitrogen (TN, DN, PON, NO₃-N) species. Proportions of particulate organic C and N were determined by difference between total and dissolved fractions.

? Under spruce, throughfall concentrations of most C and N fractions were twice as high as under beech. At both sites, concentrations and fluxes were significantly higher during the growing than the dormant season. At the broadleaved site, 80% of the annual fluxes of the DOC and TOC and 70% of the DN and TN were released during the growing season, compared to 60% for C and N at the coniferous site. POC under beech contributes with up to 30% to TOC compared to less than 20% at the spruce site.

? We suggest that pollen deposition, insect excretions and accumulated organic matter mobilised by dry/wet precipitation patterns play a supreme role for the formation of DOM and POM in forest canopies. The study demonstrates that the canopy is an important source for POM. Dynamics of DOM and POM are mainly driven by tree species effects and seasonality as well as by biotic agents.

     

Effects of a spruce budworm outbreak on element export below the rooting zone: a case study for a balsam fir forest

? Spruce budworm outbreaks are among the major natural disturbances affecting the dynamics and functioning of Canadian boreal forests. However, the element losses potentially associated with spruce budworm outbreaks have not been quantified.

? We evaluated the influence of spruce budworm outbreaks on nutrient export from boreal forest soils by comparing nutrient leaching losses during a spruce budworm outbreaks episode (1981–1984) to an unperturbed period (1999–2003) in a calibrated catchment located in a balsam fir forest.

? Nutrient soil leaching losses were significantly higher during the spruce budworm outbreaks (1981–1984) for N-NO₃ (30.1 fold), K (8.3 fold), N-NH₄ (6.2 fold), Mg (2.7 fold) and SO₄ (2.2 fold), as compared to an unperturbed period (1999–2003). When the recurrence of spruce budworm outbreaks (33 years) and a plausible average length of such events (5 years) are taken into consideration, it is estimated that in the long term, 5.6 more NO₃, 1.5 more K and 1.2 more NH₄ are leached from the soil profile during outbreaks.

? The important leaching losses during spruce budworm outbreaks, when added to the losses due to tree harvesting and fire (and acid deposition for K), may have considerable effects on soil fertility and ecosystem sustainability.

     

Effects of some ecological variables on carabid communities in native and non native forests in the Ibaizabal basin (Basque Country: Spain)

? Carabid beetles were investigated at five different forest types in the Ibaizabal basin (northern Spain). The landscape is characterized by the presence of remnants of native forest surrounded by conifer plantations.

? Carabids were trapped in 52 stands of mixed forest, beech forest, holm oak forest, mixed pine and Monterey pine plantations in 2005 and 2006. The main objectives of the study were: compare carabid diversity, recognise the characteristic species, and study the effects of ecological variables on carabid assemblages in the different forest types.

? No significative differences in species abundance, richness and diversity were found among the studied forests. Most of the trapped beetles were identified as forest generalists, nevertheless some native and non-native forest specialist species were also found. Distribution of carabid communities overlapped and, except for beech forest, no specific communities were distinguished. Altitude, percentage of grass coverage and temperature were the main variables influencing species distribution.

? The results suggest high habitat homogeneity, caused by regeneration in pine plantations of the indigenous understorey, and by poor habitat quality in native forest (patchy remnants enclosed in conifer plantations). This situation could explain the similar carabid diversity.

     

Is forest mushroom productivity driven by tree growth? Results from a thinning experiment

? Most of the edible forest mushrooms are mycorrhizal and depend on carbohydrates produced by the associated trees. Fruiting patterns of these fungi are not yet fully understood since climatic factors alone do not completely explain mushroom occurrence.

? The objective of this study was to retrospectively find out if changing tree growth following an increment thinning has influenced the diversity patterns and productivity of associated forest mushrooms in the fungus reserve La Chanéaz, Switzerland.

? The results reveal a clear temporal relationship between the thinning, the growth reaction of trees and the reaction of the fungal community, especially for the ectomycorrhizal species. The tree-ring width of the formerly suppressed beech trees and the fruit body number increased after thinning, leading to a significantly positive correlation between fruit body numbers and tree-ring width.

? Fruit body production was influenced by previous annual tree growth, the best accordance was found between fruit body production and the tree-ring width two years previously.

? The results support the hypothesis that ectomycorrhizal fruit body production must be linked with the growth of the associated host trees. Moreover, the findings indicate the importance of including mycorrhizal fungi as important players when discussing a tree as a carbon source or sink.

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Why and where do adult trees die in a young secondary temperate forest? The role of neighbourhood

? The density and identity of tree neighbourhood is a key factor to explain tree mortality in forests, especially during the stem exclusion phase.

? To understand this process, we built a logistic model for mortality in a spatially explicit context, including tree and neighbourhood predictors. Additionally, we used this model to build mortality risk frequency distributions. Finally, we tested this model against a random mortality model to predict the spatial pattern of the forest.

? Annual mortality rate was high for pedunculate oak (Quercus robur, 6.99%), moderate for birch (Betula celtiberica, 2.19%) and Pyrenean oak (Q. pyrenaica, 1.58%) and low for beech (Fagus sylvatica, 0.26%). Mortality risk models for pedunculate oak and birch included stem diameter, tree height, canopy position and neighbourhood. Mortality was affected by the specific nature of the neighbourhood showing a clear competitive hierarchy: beech > pedunculate oak > birch. Models based on random mortality and logistic regression model were able to predict the spatial pattern of survivors although logistic regression predictions were more accurate.

? Our study highlights how simple models such as the random mortality one may obscure much more complex spatial interactions.

     

Potential use of pine plantations to restore native forests in a highly fragmented river basin

? In forests, the substitution of broadleaf species by conifers can reduce biodiversity because coniferous forests generally provide less diverse vascular understories than broadleaf forests. However, in some cases, former pine plantations might be useful for restoring native forests. We compared plant species composition on the plot scale in natural beech and mixed oak forests with that in plantations of Pinus radiata. Links between plant diversity and landscape parameters (patch size, fractal dimension and distance to the nearest patch of the same type) were investigated.

? The objective of this study was to evaluate the use of pine plantations for restoring native diversity in a zone where native forests are very fragmented.

? Similar to oak forests, plant diversity in pine plantations was high, mainly due to the presence of generalist species. Some species characteristic of oak forests also appeared in pine plantations, suggesting the onset of natural forest regeneration.

? These results suggest that pine plantations could be used to promote natural regeneration of original oak forests. Moreover, residual native stands should be conserved as important sources of native species and their seeds.

     

Genetic parameters for grain angle in 28-year-old Norway spruce progeny trials and their parent seed orchard

? Sawn wood which acquires twist during drying is a substantial problem for the sawmill industry and is to a large extent caused by spiralled grain in the tree.

? In this study, the feasibility of using spiral grain angles as a selection trait in Norway spruce (Picea abies (L.) Karst) tree breeding and the extent of grain angle and its breeding values were investigated. Grain angles under bark and diameter were measured at breast height in three 28-year-old Norway spruce progeny trials and in their parent seed orchard.

? The mean grain angle value was 1.76° in the progeny trials and 1.84° in the seed orchard. Estimated genetic standard deviations for grain angle were almost 1°. Progeny trial heritabilities (H ² > 0.3) were larger than the seed orchard heritability (H ² = 0.24). Genotype by environment interaction among the progeny trials was negligible for grain angle. Progeny trial grain angles exhibited genetic correlation with seed orchard grain angles in the range 0.66 to 0.84.

? Thus, mature wood grain angles are also under considerable genetic control, raising the opportunity to reduce the occurence of left-grained trees, which are prone to producing twisted wood.

     

Genetic structure and variability of phenological forms in the European beech (Fagus sylvatica L.)

? Microsatellite markers were used to describe the genetic structure and variability of early, intermediate and late phenological forms of European beech (Fagus sylvatica L.). Two hundred and seventy individuals from three populations located in southern Poland were divided into three forms according to the phenological criterion — bud burst, and analyzed for allelic variation at five highly polymorphic microsatellite loci.

? Population differentiation was moderate and differed significantly among phenological forms. Average values of F _ST and R _ST decreased across phenological forms and amounted to F _ST values of 0.135, 0.110 and 0.108 and R _ST values of 0.365, 0.231 and 0.098 for early, intermediate and late forms of beech, respectively.

? Analysis of Molecular Variance (AMOVA) revealed different genetic structures characteristic of respective phenological forms of beech. The amount of within-population variability increased with the delay of the beginning of vegetation and amounted to 64%, 77% and 90% of total variability, depending on phenological form. A similar trend was found in average pairwise genetic distance between individuals belonging to a given phenological form (11.78, 11.85 and 12.22, from early to late forms).

? Our results demonstrate the importance of late spring frosts as a factor influencing the genetic structure of beech, and as a cause of the decrease in genetic variability as well as the increase in population differentiation proportional to the degree of phenological earliness.

     

Effect of extractives on conferred and natural durability of Cupressus lusitanica heartwood

? Identification of extractives present in Cupressus lusitanica heartwood has been conducted using GC-MS analyses. The chromatogram of toluene/ethanol extracts indicated the presence of large amounts of benzaldehyde and numerous terpenic compounds such as cedrol, agathadiol, epimanool, bornyl acetate, α-cedrene and β-cedrene.

? The effect of these extractives on the natural durability of cypress wood was investigated on heart wood blocks exposed to pure culture of Poria placenta before or after solvent extraction. Weight losses revealed severe fungal degradations on the extracted blocks compared to unextracted ones.

? Efficiency of heartwood extractives as inhibitors of the growth of Poria placenta on malt/agar test confirms their contribution to cypress natural durability.

? Cypress blocks were treated at 240 °C for different times to reach different levels of thermodegradation to evaluate effect of heat treatment on fungal durability. Results indicate that evaporation of volatile extractives during the first few minutes of heat treatment contribute to decreased wood durability, while longer treatment times lead to the expected improvement of durability.

? This study suggests that the content of extractives, which may be modified during wood drying or weathering processes, could be the origin of the conflicting data described in the literature concerning cypress natural durability.

     

Relationships of climate and cell features in stems and roots of black spruce and balsam fir

? The anatomical differences of mature black spruces and balsam firs were examined at stem and root level in order to characterize their wood properties at cellular level and link these differences to climate.

? Anatomical variability of these species was evaluated in relation to climate data gathered from 2001 to 2004 during the cell enlargement (CE) and wall thickening and lignification (WTL) phases. Lumen area, single cell wall thickness and total tracheid radial diameter were analyzed and regrouped into earlywood and latewood.

? Results from a principal component analysis (PCA) indicated that both first eigenvectors account for 82% and 90% of total variance for CE and WTL respectively. These component factors revealed that precipitation, humidity and number of days with precipitation significantly influence the lumen area (p = 0.0168) and radial cell diameter (p = 0.0222) in earlywood. Significant differences were registered between species and tree parts (stem and root) for the lumen area, radial cell diameter and cell wall thickness in both earlywood and latewood.

? In our study, black spruce exhibited smaller tracheid size in both stem and roots compared to balsam fir. Furthermore, the lower amount of tracheids produced during the growing season and higher proportion of latewood ensure a higher wood density of black spruce. The influence of temperature on earlywood formation is significant, whereas no influence was observed on latewood.

     

Estimating growth in beech forests: a study based on long term experiments in Switzerland

? This contribution presents a dynamic stand growth model for Beech (Fagus sylvatica L.) forests, based on a dataset provided by the Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Birmensdorf. The dataset includes 143 research plots, covering a wide range of growing sites and providing up to 16 interval measurements per research plot.

? The objective of this research is to complement the range of existing beech growth models by bridging the gap between the historical yield tables and the single tree growth models. The specific aim is to develop transition functions which will project three state variables (dominant height, basal area and number of trees per hectare) at any particular time, in response to any arbitrary silvicultural treatment.

? Two of the transition functions were derived using the generalized algebraic difference approach (GADA), the third one was derived with the algebraic difference approach (ADA). All the functions were fitted simultaneously using iterative seemingly unrelated regression and a base-age-invariant method. The influence of thinnings on basal area growth was included by fitting different transition functions for thinned and unthinned stands.

? The overall model provides satisfactory predictions for time intervals up to 20 years. The new model is robust and its relatively simple structure makes it suitable for economic analysis and decision support.

     

Comparison of methods to quantify loss of hydraulic conductivity in Norway spruce

? Percent loss of hydraulic conductivity (PLC) is an important measure in plant water relations, but the available methods differ and results have rarely been compared.

? We compared PLC measured in Norway spruce (Picea abies) by quantifying conductivity before and after removing emboli, either by flushing with high water pressure or by infiltration under a partial vacuum, with relative water loss and staining conductive xylem to test advantages and possible problems of commonly applied methods.

? Infiltration removed nearly all gas from sections of drought-stressed and unstressed twigs, and flushing and infiltration efficiently removed emboli. Infiltration appears less prone to producing clogged xylem elements than high pressure flushing. Relative water loss (RWL) and the proportion of xylem stained with phloxine B (PSX) was also highly correlated with PLC, the latter can be quantified by image analysis, and also shows the pattern of xylem dysfunction. Loss of conductivity was more common in the inner annual rings, in early wood within an annual ring, and in compression wood, though pattern differed substantially between branches.

? Advantages and potential problems of these methods are discussed and it is suggested that RWL or PSX may be used as proxy measures for PLC in species when the correlations have been established.

     

Comparing needle and shoot phenology with xylem development on three conifer species in Italy

? Since the 1960s biologists have been trying to assess factors affecting cambial activity and latewood formation. A comparison of the phenology of the apical and lateral meristems could add new clues in order to understand mechanisms of the growth dynamic in conifers.

? This study compared needle and shoot growth with xylogenesis in Larix decidua, Pinus cembra and Picea abies during 2001 to verify if cambial resumption occurred after shoot and needle growth had begun, and if latewood initiation occurred when shoot lengthening was complete.

? In pine and spruce, needle and shoot lengthening was synchronous between mid-June and August while larches showed an early development of the needles at the end of May with a later shoot lengthening. Cell production lasted from mid-May to the beginning of August, with the first differentiating cells observed at the end of May, which indicated a earlier reactivation of the cambium with respect to needle and shoot growth.The first latewood cells began wall thickening between July and August, during the termination of needle and shoot lengthening.

? The hypothesis that cambial resumption occurs after shoot and needle elongation was rejected. The separation in time between apical growth and secondary wall thickening of latewood suggested the presence of an internal competition in resource allocation.