Heartwood and sapwood allometry of seven Chinese temperate tree species

? Allometry of sapwood/heartwood is essential for understanding tree growth, water transport and carbon allocation, timber production and use, but such an allometry is lacking for Chinese temperate tree species.

? We studied the allometry and development of heartwood and sapwood for seven Chinese temperate tree species: Korean pine (Pinus koraiensis Sieb. et Zucc), Dahurian larch (Larix gmelinii Rupr.), Japanese elm (Ulmus davidiana Planch var. japonica (Rehd.) Nakai), Manchurian ash (Fraxinus mandshurica Rupr.), Manchurian walnut (Juglans mandshurica Maxim.), Amur cork-tree (Phellodendron amurense Rupr.), and Mongolian oak (Quercus mongolica Fisch.).

? All heartwood parameters investigated, including heartwood radius (HR), heartwood formation rate (HFR), heartwood ring number (HRN), heartwood initiation age (HIA), and heartwood volume ratio (HVR), were positively correlated with tree cambial age (CA). The HR, sapwood width (SW), sapwood area (SA), heartwood and sapwood volumes were significantly related to stem diameter at breast height (DBH) or xylem diameter. There was a polynomial relationship between the sapwood ring longevity (SRL) and sapwood ring number (SRN). However, most of the allometric relationships were species-dependent. The hardwood formation patterns were different between coniferous and broadleaved tree species. A power function was suitable to scale SA from DBH, but the exponent varied from 1.32 for the larch to 2.19 for the cork-tree.

? Our allometry provided a practical means to assess wood development and related physiology for the temperate tree species.

     

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.

     

Extending sapwood — Leaf area relationships from stems to roots in Coast Douglas-fir

? Studies of allometric relationships between leaf area and the cross-sectional area (CSA) of sapwood in the stem have shed light on the structural and functional relationships between water-conducting and photosynthetic tissues.

? The purpose of this study was to test whether sapwood-leaf area relationships could be extended from stems to roots in coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Twelve trees were felled, their stumps were excavated, and the CSA of sapwood and heartwood were estimated for individual roots, entire root systems, and stem section.

? Root sapwood CSA was greater than sapwood CSA throughout the stem, and the ratio of leaf area to sapwood CSA (A _l :A _s ) was accordingly lower for root sapwood. The relationship between sapwood CSA and leaf area was more variable in roots and at groundline compared to crown base. Root A _l :A _s decreased with relative tree height (tree height/mean stand height).

? The strong allometric relationship between leaf area and the CSA of sapwood in the stem generally holds when extended to roots. The greater CSA of sapwood in roots versus stems may reflect differences in their roles in supporting the tree.

     

Improving models of forest nutrient export with equations that predict the nutrient concentration of tree compartments

? The objective of this study was to explore the distribution of major nutrients (N, P, K, Ca and Mg) in the aboveground compartments of an intensively managed tree species (Pinus pinaster Ait.). A total of 53 trees were cut down in even-aged stands respectively 8, 16, 26, 32 and 40 years old. The nutrient concentrations of the aboveground compartments were analysed.

? Nutrient concentrations of foliage did not vary with any of the variables used, except needle age. Nutrient concentrations of living branches, stem bark, stem sapwood, stem heartwood, stemwood and stem decreased with increasing branch diameter, bark thickness, sapwood thickness and heartwood thickness, respectively. Beyond a certain value of the predictive variable (stem diameter ≈ 15 cm; branch diameter ≈ 2.5 cm), the concentration of all the nutrients stabilised.

? A 50 year-old pine stand was used to obtain a validation dataset for nitrogen concentration. For this nutrient, the regression relationships gave satisfactory estimates for most compartments (mean error = 12–25%) and particularly for the stem.

? A procedure is proposed to estimate the nutrient exports associated with harvests of Pinus pinaster biomass.

     

Colour and decay resistance and its relationships in Eperua grandiflora

? Eperua grandiflora, which is widely distributed in the French Guiana forest region, shows high variability in decay resistance. Further information concerning this wood quality parameter is necessary, but standard testing methods are complex and time-consuming. We assessed the use of colorimetry to determine durability in heartwood samples from a range of trees.

? Eperua grandiflora colour parameters were measured using a CIELAB system, revealing that the tree effect was greater than the radial position and height effects.

? The wood samples were exposed to Coriolus versicolor and Antrodia sp. according to two European standards (En 350-1 and XP CEN TS 15083-1). Eperua grandiflora is more susceptible to brown rot. These two standards did not give the same durability classes. The high variation in natural durability was due to the tree effect.

? These two properties were found to be correlated and the assessment also distinguished the extreme durability classes but they are not sufficient to classify the class of durability of this species.

     

Stimulation of tree defenses by Ophiostomatoid fungi can explain attack success of bark beetles on conifers

? Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.

? After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.

? We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.

1. Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.
2. We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.

? We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).

? Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:

1. any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;
2. the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;
3. Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.

? We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.

? We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.

     

Fecundity and larval development of Monochamus galloprovincialis (Coleoptera Cerambycidae) in experimental breeding

?The study aimed at clarifying basic life traits (fecundity, mortality and development) of Monochamus galloprovincialis, the vector of Bursaphelenchus xylophilus, the causative agent of the Pine Wilt Disease, in Portugal.

? Mating and oviposition experiments were conducted on P. sylvestris logs under laboratory conditions during two years. Larval development was followed outdoors.

? Fecundity was high, with 138.2 eggs per female.

? For the first time, the number of larval instars (4) in M. galloprovincialis was recognized and instars were described.

? Informations are provided on the shape and size of the larval galleries. Boring of galleries in sapwood and heartwood started at the third instar. All instars could overwinter but adults emerged simultaneously resulting in a univoltine life cycle for the majority of the individuals.

? An obligatory diapause in the forth instar was assumed. However, 8.1% of the insects had a two year development.

? Important differences in mortality were noted between the two experimental years, which could be explained by differences on the size of the breeding logs.

? There is a high biological proximity between M. galloprovincialis populations in France and those in Portugal and Northern Europe. However, a higher fecundity and longevity was observed in France, as well as a higher percentage of insects with a two year development, compared with portuguese populations.

     

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.

     

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.

     

Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

Colour and chemical changes on photodegraded beech wood with or without red heartwood

The focus of this study was to investigate the chemical and colour changes occurring at different exposure times on artificially photo-irradiated surfaces of normal and red heartwood in beech in order to understand the mechanisms that cause the changes and to evaluate the possibility of usages of beech not only for energy production purposes. In this sense, surface colour modifications are of crucial importance to define the commercial value of beech wood. The artificial photo-irradiation of the wood samples was performed in a Solar Box, equipped with an ultraviolet filter that cuts off the spectrum at 280 nm. Reflectance spectrophotometry and Fourier transform infrared (FTIR) spectroscopy were used to assess artificial sunlight influence. The experimental data were statistically treated to evaluate their significance. Colour monitoring revealed that wood surface colour undergoes an important variation due to photo-irradiation, occurring within the first 24–48 h. Moreover, it was found that the chromatic coordinates (L*a*b*) in normal wood and in red heartwood tended to similar values after 504 h. FTIR spectroscopy allowed for investigating the rate of photodegradation of wood surface due to oxidation reactions of wood components. The results were validated by statistical analysis applied both to the colorimetric and spectroscopic data.     

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.

     

Radial distribution of quinones in plantation teak (Tectona grandis L.f.)

? Quinones are primarily responsible for the natural durability of teak.

? The previous paper in this series reported on the natural termite resistance of teak trees of different ages (8-, 30- and 51-year-old trees). In this study, the radial distribution of quinones (tectoquinone, lapachol, desoxylapachol and its isomer) and other components in the ethanol-benzene (1:2) extract were measured by means of gas chromatography.

? Significant differences in desoxylapachol or its isomer content were found among the outer heartwood of 8-, 30- and 51-year old trees, as well as between the inner and outer parts of the heartwood.

? All toxic quinone contents were positively correlated with the total extractive content. The highest correlation degree was measured in the isodesoxylapachol content.

? Although linearly related, only modest correlations were observed between the natural termite resistance parameters and the content of tectoquinone and isodesoxylapachol.

     

Variation of wood color parameters of Tectona grandis and its relationship with physical environmental factors

? Context

Teak??s wood color is considered an important attribute in the marketing phase and it has been influenced by environmental setting, stand conditions and management, plant genetic source, and age. However, there is a lack of understanding about how the environmental factors might affect the teak??s wood color planted in short-rotation forest plantations.

? Aims

The aim of this study is to understand the relationship, gathered from generated information, between edaphic and climatic variables and their effects in the wood color variation of Tectona grandis from trees in forest plantations.

? Methods

Twenty-two plots were grouped in five cluster sites that shared similar climatic and soil conditions. Data about soil??s physical?Cchemical properties and climatic variables were collected and analyzed. Representative trees were harvested next to each plot in order to obtain a wood sample per tree at a diameter breast height. Wood color was measured using standardized CIELab??s chromaticity system.

? Results

After comparing the wood change color index (?E*) in the five studied clusters, it was found that heartwood produced from drier and fertile sites had more yellowish-brown color. The heartwood b* color index resulted with significant correlations (R?>?0.5, P?<?0.05) among nine climatic and eight edaphic variables.

? Conclusion

It was concluded that climatic variables should be considered as the first-order causal variables to explain wood color variation. Hence, darker b* wood color was associated with dry climates; also, with deeper and fertile sites.     

Variation in carbon concentration and basic density along stems of sessile oak (Quercus petraea (Matt.) Liebl.) and Pyrenean oak (Quercus pyrenaica Willd.) in the Cantabrian Range (NW Spain)

Context

Information about variations in basic density (BD) and carbon content (%C) along tree stems is key to assessing forest carbon sinks.

Aims

The aim of the study was to determine any differences in %C and BD between different woody tissues (bark, sapwood and heartwood) in two widespread European oak species (Quercus pyrenaica and Quercus petraea).

Methods

Twenty trees were felled in northern Spain, and 317 discs cut from the trees were dried and analysed to determine %C and BD.

Results

There were significant differences in %C between bark, heartwood and sapwood, and between species. There were also significant differences in BD between the tissues (heartwood?>?sapwood?>?bark), and the BD was higher in Q. petraea than in Q. pyrenaica. Both %C and BD varied along the stem.

Conclusion

More accurate estimates of carbon contents were obtained by using specific values for different heights and anatomical parts, than by using single values.     

Disentangling the effect of size and competition: a growth model for Aucoumea klaineana

• Context Okoume (Aucoumea klaineana) is the most important timber species in Gabon, while being classified as vulnerable in the IUCN red list. Thousands of growth measurements for this species have been performed since the 1950s but, because of a lack of integrated analysis, did not bring a consistent view on its growth pattern.
• Aims This study aims at disentangling the effects of tree size and local competition on tree growth.
• Methods A growth model was fitted for okoume, using data from seven sites in Gabon and Congo. The growth model was designed to be useful for forest management, which means that it relied on variables that could be computed using forest inventory data.
• Results A lognormal growth model with a negative response to stand basal area was selected. A significant residual site effect on growth was found, with a slower growth in the sites near the border of the natural range of okoume than in the sites far from it.
• Conclusion Growth strongly responded to local stand attributes such as tree density and basal area. Growth decrease with stand age in monodominant okoume stands was correctly predicted, although tree age was not incorporated into the model.

     

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.

     

Use of wood shrinkage characteristics in breeding of fast-grown Acacia auriculiformis A. Cunn. ex Benth in Vietnam

u| li]? Genotypic variation in wood total and partial shrinkage, basic density and growth traits was estimated in 51÷2 year old Acacia auriculiformis trees in a clonal test. li]? In the tangential, radial and longitudinal directions, the mean values were 2.64%, 1.64% and 0.77% for partial shrinkage, and 5.92%, 3.23%, and 0.96% for total shrinkage, respectively. Total and partial transverse shrinkage were significantly greater in sapwood than in heartwood. li]? Clonal repeatability (H _C ² ) estimates for partial shrinkage were lower than those for total shrinkage, and heartwood shrinkages had lower H _C ² than those for sapwood. Estimates of H _C ² were from 0.32 to 0.38 for total transverse shrinkage, comparable to H _C ² for both total and partial volumetric shrinkages (0.40 and 0.32, respectively). However, H _C ² for longitudinal shrinkages, total and partial coefficients of anisotropy were only from 0.09 to 0.18. li]? The genotypic coefficients of variation of all shrinkage traits varied from 5.45% to 8.02%. Total shrinkage was strongly positively correlated with partial shrinkage in each dimension. Genotypic correlations were not significant between shrinkage and growth traits or density. Strong and significant correlations between transverse shrinkage in heartwood and sapwood (0.87) indicated that juvenile shrinkage is a good genetic indicator of this trait in older trees. Utilisation du retrait du bois dans l’amélioration de l’Acacia auriculiformis A. Cunn. ex Benth à croissance rapide au Vietnam.     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

Growth response of Pinus pinaster Ait. to climatic variables in central Spanish forests

? The effect of climatic variables (temperature and precipitation) on radial growth of the Mediterranean Maritime pine (Pinus pinaster Ait.) was studied using dendrochronological techniques in the Iberian Peninsula.

? Ten tree-ring width chronologies, along the central distribution area of the species, were built. Chronology variability was analysed using Principal Component Analysis (PCA) for the period 1952–2005.

? The first principal component (PC1) explained 56% of tree-growth variability. Tree-growth association with climate was analysed at regional and local scales using correlation coefficient and bootstrapped response functions.

? Radial growth at both scales was positively correlated with rainfall during and prior to the growing season at all sites, and with summer rainfall before the growing season at five sites. Mean temperature effect changed according to the sampling site, from non-significant at the highest sites to significant (positive relationship in winter) at the lowest sites. Growth season temperature also had a negative effect.

? The Kalman filter was applied to estimate changing association between growth and climate over-time. Results suggested a change in association, initiated in the 80s, from non-significant to significant (*p < 0.05) at six of the sampling sites.

? Pinus pinaster is an accurate species for analysing tree-growth association with climate and for studying plant behaviour under global change conditions.