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.

     

Effects of the lateral growth rate on wood quality of Gmelina arborea from 3.5-, 7- and 12-year-old plantations

? Awareness of the shortage of fossil resources leads to an increasing demand for woody biomass. We investigated the feasibility of using fast-growing Gmelina arborea wood for material production. Gmelina arborea wood samples were collected from trees of varying cambium ages in Indonesia, from 3.5-, 7- and 12-year-old plantations.

? The lateral growth rate and the cambium age did not significantly affect the longitudinal released strain of the growth stress, xylem density, or microfibril angle at the outermost surface of the secondary xylem at any sampling site. However, fiber length in the 3.5-year-old plantation tended to be shorter in smaller diameter trees, whereas in larger diameter trees it was almost the same as that in trees from the 7- and 12-year-old plantations. This suggests that smaller diameter trees in the 3.5-year-old plantation had not yet produced mature wood.

? Xylem qualities had already reached values appropriate for harvesting, except in the smaller diameter trees from the 3.5-year-old plantation. This indicates that the larger diameter trees had already matured, regardless of their cambium age. These results suggest that the next step is to develop silvicultural treatments to increase the lateral growth rate during the early growing stage, in order to produce as much mature wood as possible, as quickly as possible.

     

High resolution temporal variation in wood properties in irrigated and nonirrigated Eucalyptus globulus

? Environmental determinants of wood properties variation were examined in Eucalyptus globulus, a globally important hardwood plantation species, in southern Tasmania, Australia.

? Radial variation in wood properties, measured with the SilviScan system, were re-scaled from distance to time abscissa using stem radial growth data measured with dendrometers. With this re-scaled data it was possible to evaluate how water availability and temperature affected wood density, microfibril angle (MFA) and fibre and vessel transverse dimensions in irrigated and non-irrigated trees.

? Wood density, fibre radial diameter and MFA were sensitive to water availability. Wood density increased and fibre radial diameter decreased in response to reduced water availability. When high water availability was maintained, wood density was negatively correlated with temperature. Together, temperature and soil matric potential explained about 60% of temporal variation in wood density variation. In contrast MFA was not related to temperature but decreased with increasing water stress. Slower growing trees also had lower MFA than faster growing trees. Slower growing trees had a larger number of vessels per unit area of wood than faster growing trees within this even aged stand. However, vessel radius to the 4th power was significantly higher in faster growing trees than in slower growing trees.

? Overall, E. globulus wood properties were sensitive to temporal changes in environmental conditions (particularly water availability) and associated growth rates. The data provided support for the hypothesis that growth rates are hydraulically mediated.

     

Physical properties of wood in thinned Scots pines (Pinus sylvestris L.) from plantations in northern Spain

? The physical properties of wood and the associated variations within and between trees were evaluated by analysing 770 small specimens of clear wood from 11 Pinus sylvestris L. trees thinned from 3 plantations.

? Within-tree variations in basic density or volumetric shrinkage increased with cambial age and decreased with increasing ring width. The effect of the height in the stem on wood properties was considered indirect and height was not included as an explanatory variable in the mixed models proposed to estimate basic density and volumetric shrinkage.

? The models had random components for the intercept parameter and explained 52.5% of the total variance in basic density and 56% of the total variance in volumetric shrinkage. Linear shrinkage in the direction of the grain was extremely variable.

? Between-tree variation and between-plot variation in the physical properties of wood were high, considering that all trees sampled were growing in similar sites and stands. It would then be desirable to predict physical properties of wood on living trees in order to use the quality of wood as a criterion for timber tree selection in thinnings.

     

Variation in the natural termite resistance of teak (Tectona grandis Linn. fil.) wood as a function of tree age

? Tree age is one of the most important factors to affect the natural durability of wood.

? The purpose of this study was to determine the natural termite resistance of heartwood and sapwood of teak (Tectona grandis Linn. fil.) for trees aged 8, 30 and 51 years. Reticulitermes speratus Kolbe was employed as a test termite using a no-choice feeding method.

? The heartwood and sapwood of all of the trees tested exhibited antitermitic activity. Based on the mean mass loss due to termite activity, the sapwood and heartwood regions of 8-year-old trees are the most susceptible to termites while the heartwood regions of the 30-year-old trees showed termite resistance similar to the termite resistance of 51-year-old trees.

? The mass loss is moderately correlated with n-hexane extractive content, total extractive content, brightness and redness of the wood.

     

Integration of water transport pathways in a maple tree: responses of sap flow to branch severing

? It has been known for a long time that sectored and integrated patterns of vascular systems exist in different species and even within the same tree, depending on its age and history. However, very few publications consider the topology of the vascular pathways between roots and branches.

? Some results on this important aspect of the vascular system are presented in this paper. They have been obtained with adult maple trees by directly studying the water movement in the stem and root xylem with the heat field deformation (HFD) method for sap flow measurements.

? Multi-point HFD sensors were installed at different heights of a Norway maple tree (Acer platanoides L.) along its stem axis. Single-point HFD sensors were installed in three small lateral roots of another sample maple. Experimental treatments (branch severing) triggered changes in sap movement in the stem and root sapwood.

? The sample trees belong to the group with an integrated transport system (“integrated pipes”), sharing stem space on both sides of the tree to supply two large parts of the crown with water from each root sector. Nevertheless, conducting pathways had their autonomy for axial transport and the pipe model theory describes the vascular system of the studied trees well. Thus, the integration of axial transport in the stem xylem should presumably occur through the cross-grained network of axial vessels.

     

Ranking larch genotypes with the Rigidimeter: relationships between modulus of elasticity of standing trees and of sawn timber

? Direct assessment of modulus of elasticity (MOE) on standing trees is attractive for breeders to evaluate genotypes prior to selection: this can be done using the Rigidimeter, a bending-based measurement device.

? In this study, we tested its reliability to properly rank genotypes by relating trunk MOE with MOEs estimated with a vibrating analysis system (Bing) on different types of conditioned wood specimens from the same trees (boards and standardised 2×2×30 cm-clear-wood specimens). One hundred and ten trees from different genotypes of hybrid larch (Larix × eurolepis) were tested.

? Mean trunk MOE was 7 300 MPa with a similar value obtained for sawn boards. Clear-wood specimens MOE increased from pith to bark from less than 6 000 MPa to nearly 9 000 MPa. Moderate correlations (r = 0.48–0.61) were found at the individual tree level between trunk MOE and MOE of wood samples.

? Single specimen MOE was shown to be strongly related to a linear combination of trunk MOE and sample position.

? At the genotype mean level, trunk MOE was highly correlated with wood samples MOE (r = 0.80–0.91). Ranking of genotypes based on trunk MOE was mostly consistent with that based on standardised specimens.

? It was concluded that besides other operational advantages which are discussed, the Rigidimeter is a valuable tool for breeders to routinely evaluate and rank genotypes for stiffness prior to further selection.

     

Variation in the decay resistance of larch to fungi

? Decay resistance of larch (Larix sp.) to fungi was evaluated on heartwood samples belonging to 3 species (L. decidua, L. kaempferi and their hybrid), 3 races of European larch (polonica, sudetica and alpine), 13 wood lots (populations) and 313 trees.

? Larch wood appeared, on average, as moderately durable although a high variability was observed. At the sample level as well as at the mean individual tree level, durability ranged from class 1 to 5 according to EN 350-1 standard. At the population level, larch wood varied from ‘durable’ to ‘slightly durable’. Genetics played a major role in decay resistance at the species, provenance and tree levels. Environmental factors such as the position of heartwood samples and the age of trees were also identified as a source of variability.

? The most durable wood was not necessarily from old native alpine stands of European larch: some young larches from faster growing lowland origins also produced durable wood.

? Genetic improvement of larch wood durability appeared therefore likely by the selection of the best populations for decay resistance as well as from the selection of individuals.

     

Xylem traits in European beech (Fagus sylvatica L.) display a large plasticity in response to canopy release

Key message

The position of trees in the canopy impacts xylem structure and its inter-annual variation. After canopy release, the increase in the hydraulic conductivity of growth rings was driven by an increase in radial growth in large trees, and by both an increase in radial growth and changes in xylem structure in saplings.

Context

Forest canopies are frequently subjected to disturbances that allow understory trees to access the upper canopy. The effect of canopy release on xylem anatomy has been assessed in juvenile trees and saplings, while the potential acclimation of larger trees remains poorly documented.

Aims

We estimated the potential hydraulic conductivity of growth rings in large understory trees compared to overstory trees, and evaluated the responses to canopy release in large trees and in saplings.

Methods

We recorded radial growth, wood density, and vessel structure in beech trees according to their position within the canopy and their size. Xylem traits were followed during 6 years after canopy release for large trees, and during 2 years for saplings. Vessel diameter and frequency as well as ring area were used to compute the potential annual ring hydraulic conductivity.

Results

Large understory trees displayed lower radial growth increments and lower potential annual ring hydraulic conductivity than overstory trees. After canopy release, potential annual ring hydraulic conductivity increased in large trees, due exclusively to increased radial growth without any change in specific hydraulic conductivity. It increased in saplings due to both increased radial growth and increased specific conductivity.

Conclusion

Tree size impacted xylem structure and resulted in plasticity of the potential hydraulic conductivity of the annual tree ring following canopy release.

     

Effects of site on fibre, kraft pulp and handsheet properties of Eucalyptus globulus

? Eight-year old trees from two Eucalyptus globulus Labill. clones planted across three different sites in Tasmania, Australia, were sampled for wood and kraft pulp/handsheet properties.

? Site had a significant effect on all measured properties. Compared with the poor site (Parkham) the wood from the good site (West Ridgley) had on average 11 % lower wood density. The poor site had also greater microfibril angles, shorter fibres at lower pulp yields.

? The handsheets produced with pulp from the poor site resulted in comparatively higher bulkiness, lower burst, lower tear and tensile indices, lower zero span tensile strength, but higher opacity, higher light scattering and higher surface roughness. Significant height effects were found with all wood properties, and also with tear index, zero span tensile strength and opacity.

? Discriminant analysis showed that for 76 out of 100 handsheets the raw material source, i.e. growth site, could be predicted correctly using a set of handsheet properties with tear index and bulk index being most prominent.

? This is unique evidence that site conditions are strongly reflected in handsheet properties produced from Eucalyptus pulp.

     

Genetic variation in wood density and correlations with tree growth in Prosopis africana from Burkina Faso and Niger

? Variation in wood basic density and its correlation with tree growth were investigated at 13 years in a provenance/progeny test of Prosopis africana in Niger. The test included progeny from 256 trees sampled from 24 provenances in the Sahelian ecozone of Burkina Faso and Niger.

? Variation in wood density was significant due to provenances and families within provenances. Individual tree heritability was higher for wood density than for growth traits. Provenance means for wood density increased from the more humid to the drier parts of the sample region. Phenotypic correlations indicated that larger trees tended to have denser wood.

? Clines suggest that natural populations of P. africana from the drier parts of the sample region have the genetic capacity to produce denser wood, compared with populations from the more humid parts of the region. Correlations and heritability estimates suggest that selection of faster-growing trees may produce a small gain in wood volume but little (if any) gain in density in the subsequent generation. Multi-location provenance/tests are needed to confirm this tentative conclusion.

     

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.

     

Tree mortality in a mixed deciduous forest in Northwestern Russia over 22 years

? We studied mortality rates of birch, small-leaved linden, pedunculate oak, Norway maple, black alder, common ash and Norway spruce, assessed through two inventories of 12 664 trees in the unmanaged mixed deciduous forests of Oranienbaum Park, northwestern Russia, in 1981 and 2003.

? Our objectives were to evaluate if (a) long-term mortality rates vary among species, and if (b) crown condition, age and tree size affect the probability of mortality.

? Over this period, tree mortality rates in the park varied annually from 1 to 3% for healthy and moderately healthy trees, and from 3.9 to 9.3% for declining trees. The lowest mortality rates were observed for small-leaved linden and oak (0.8 and 1.0%, respectively), and the highest rate for birch (2.7%). We found (1) a significant and consistent association between tree condition and the probability of mortality, and (2) significantly higher mortality rates in smaller trees of ash, maple and oak.

? The effect of species-specific biology on mortality rates in the Oranienbaum Park was largely overridden by the “health status” and environmental conditions of the trees (e.g. degree of crown shading). The observed rates were similar to the estimates from natural temperate deciduous forests in both Europe and North America, indicating similar intensity of mortality in these ecosystems under natural conditions.

     

Methodology to assess both the efficacy and ecotoxicology of preservative-treated and modified wood

? Wood used in outdoor conditions out of ground contact is susceptible to weathering, inducing both fungal decay and leaching of components to the environment.

? This paper presents a methodology to determine these two parameters for untreated, preservative-treated and modified wood. Therefore, the wood was first leached and subsequently exposed to fungal decay of the most prominent wood-rotting fungi. The crustacean Daphnia magna was exposed to the leachates to provide information on their impact on the environment.

? Combining both parameters reveals that preservative-treated wood and modified wood are capable of protecting the wood adequately for application under use class 3 conditions without posing a threat to the environment.

? This proves the suitability of the concept of combining efficacy and ecotoxicology for the evaluation of new types of wood treatments.

     

Non-destructive monitoring of early stages of white rot by Trametes versicolor in Fraxinus excelsior

? Non-destructive detection of fungal decay in living trees is relevant for forest management of valuable species, hazard tree assessment, and research in forest pathology. A variety of tomographic methods, based on stress wave timing, radioactive radiation, or electrical resistivity have been used to detect decay in standing trees non-destructively. But apart from mobile gamma ray computed tomography (Habermehl and Ridder, 1993) which is virtually unavailable, the detection of incipient stages of decay is still not possible.

? Wood moisture and electrolyte content influence the electric resistivity of wood. Both are changed by fungal decay. Therefore electric resistivity tomography (ERT) should detect decay in its early stages. Then it could be used to monitor the spatial and temporal progress of degradation.

? We infected four Fraxinus excelsior trees with Trametes versicolor using wooden dowels and measured two-dimensional electric resistivity tomograms 3, 10, 13 and 21 months after infection. Immediately after the last electric resistivity measurement trees were felled for further analyses of stem cross-sections. Wood moisture content and raw density had significantly increased in infected areas, but dry density had not significantly changed after 21 months. Areas of very low electric resistivity around the infected wounds correlated very well with infected wood in the stem cross-sections. Increasing areas of low electric resistivity around the infected wounds during consecutive measurements indicate increasing areas of infected wood.

? We conclude that the growth of white rot by Trametes versicolor can be monitored with electric resistivity tomography (ERT) beginning from incipient stages, even before wood density decreases. ERT could therefore be a powerful research tool for decay dynamics as well as a method for diagnosing wood decay in forestry and arboriculture.

     

Genetic parameters for early wood and latewood densities and development with increasing age in Scots pine

? Each annual ring in pines consists of earlywood and latewood with considerable difference in density and width. To get a better determination of the genetic regulation of total wood density in Scots pine (Pinus sylvestris L.), density and width of those ring sections were measured in annual ring numbers 12 to 21 of Scots pines in a full-sib progeny test. Tree height and stem diameter were also measured.

? Heritabilities for the annual ring sections increased with age for earlywood density from 0.08 to approximately 0.25; latewood density showed similar reductions. Heritability over all 10 annual rings was 0.25 for earlywood density, 0.22 for latewood density, 0.29 for height and 0.10 for stem diameter. Genetic correlations between earlywood and latewood density and growth traits were negative, while they were strongly positive between densities of adjacent annual rings (0.70–1.0).

? Despite the higher heritability of earlywood density, the strong positive genetic correlation between those traits indicates little benefit from focusing solely on earlywood density when selecting for wood density. Analysing earlywood and latewood separately does not benefit from including the width of the corresponding ring section as a covariate. Juvenile wood may possibly turn into mature wood 15–20 y from the pith.

     

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.

     

Effects of genetic entry and competition by neighbouring trees on growth and wood properties of cloned Norway spruce (Picea abies)

? The aim of this work was to study both the effects of genetic entry and competition by neighbouring trees on growth, yield, wood density traits and fibre properties of 20 Norway spruce clones grown in an experimental trial located in southern Finland. The material included 10 Finnish clones, 2 Russian clones and 8 provenance-hybrids clones, the latter ones representing crosses between Finnish and foreign parent trees.

? We found that, compared to growth and yield, wood density traits and fibre properties showed, on average, lower phenotypic variations. Moreover, significant differences could be observed among the clones regardless of the trait (p < 0.05). Conversely, on average, no clear differences could be found between Finnish, Russian and provenance-hybrids clones in most of the studied traits.

? The marked differences among the clones, and lack of clear differences among the provenances suggested that any ranking, regarding different traits, should be based on individual clones. The phenotypic correlations between different traits ranged from moderate to strong suggesting that selection based on one trait would affect other traits. Regardless of clone, the growth of trees (e.g. breast height diameter) decreased and the wood density increased with increasing competition by neighbouring trees, which was expressed by competition index.

     

Lifespan and mortality of old oaks — combining empirical and modelling approaches to support their management in Southern Sweden

? Old oaks (Quercus robur L.) play an important role in the southern Scandinavian landscape by providing habitat for a wide range of species, a large proportion of them being currently on the National Redlists.

? To provide support for the management of these trees, we review data on oak mortality and formulate a mortality-driven stochastic model analysing interactions between mortality rate, oak recruitment rate into 100–150 age class, and amount of oaks older than 200 years.

? Empirical annual mortality rates varied between 0 and 13% with average 1.68%. Trees older 200 years had an average mortality rate of 1.1%. Oaks in the high density forests showed higher mortality (3.2%) as compared to the trees growing in the low density forests (1.2%). A 400-year long modelling exercises indicated that under current mortality rates (regular mortality being centred around 1% annually; and irregular mortality 7% with average return time of 13 years) the long-term maintenance of 20 trees older than 200 years per ha would require an input rate of 1 to 5 trees × year^?1 × ha^?1 into the 100–150 years old class.

? The modelling highlighted the importance of initial oak abundance affecting amount of old trees at the end of shorter (100 years) simulation period.

     

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.