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.

     

Effects of organic residue management and legume cover on growth of pine seedlings, nutrient leaching and soil properties

? The short-term effect of organic residue management on the growth and nutrition of Pinus pinaster Ait. seedlings, and on nutrient leaching and chemical properties of an acid soil was assessed through a lysimeter experiment. Treatments included absence, placement on the soil surface, and incorporation into the soil (with and without legume cover cropping) of organic residues (forest floor litter or forest floor litter plus harvest residues).

? Residues placed on the soil surface enhanced seedling growth. Organic residues reduced nutrient losses (NO ₃ ^? , Ca and Mg) and resulted in nutrient accumulation in the soil. Harvest residues positively affected K seedling nutrition status and enhanced K soil accumulation.

? Legume cover cropping reduced soil nutrient losses (N, Ca, Mg and K) during the early stage of seedling growth; it also improved seedling nutrition status (N and P), but without any effect on growth.

? Harvest residues plus forest floor litter placed on the soil surface was the most appropriate management to both reduce nutrient losses through leaching and increase height of seedlings at the end of the experimental period (two years).

     

Wood colour variation in sapwood and heartwood of young trees of Tectona grandis and its relationship with plantation characteristics, site, and decay resistance

? Wood colour of Tectona grandis produced from fast-growth plantations is highly variable and the causes of this variation are relatively unknown.

? With the purpose of understanding the colour variation, different fast-growth plantations were sampled with different growth rates, tree ages, and sites.

? Wood colour was measured with a CIELab system, where three variables are estimated: coordinate L* for lightness, coordinate a* defines redness and coordinate b* defines yellowness.

? Results showed only a negative correlation between L* and a*. L* and a* were negatively and positively respectively correlated with pith distance in heartwood, but not for b*. No correlations were found between L* and b* in sapwood and plantation characteristics, while a* was positively correlated with age and height of tree and growth rate. In heartwood, tree age and diameter at breast height were correlated with all colour parameters, but tree height and plantation density were correlated with a* and b*. Cluster site had correlation with L*. Multiple correlation analysis showed that the heartwood is increasing darker (L*) and redder (a*) when the trees are older and bigger. Correlation coefficient shown that sapwood and heartwood with lighter colour (L*) is less resistance to fungal attack, but redness colour (a*) increasing decay resistance.

     

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.

     

Intraspecific variation in growth and allocation patterns in seedlings of Pinus pinaster Ait. submitted to contrasting watering regimes: can water availability explain regional variation?

? Interpopulation variation in key functional traits of Pinus pinaster Ait. is well recognized. However, the relative importance of drought tolerance to explain this regional variation in the species remains elusive.

? Here, we raise the question whether water availability constitutes a likely driver of regional variation in biomass allocation, growth and morphological traits of ten populations that cover the distribution range of P. pinaster. We carried out an experiment where seedlings of five families per population were submitted to two contrasting watering treatments.

? The effects of water availability and population were significant for relative diameter and height growth rate, biomass allocation and number of lateral stems and dwarf shoots. Total dry mass significantly differed between watering treatments but it did not among populations. Populations could be clustered into four main groups. Root mass fraction explained most of the variation and significantly correlated to altitude but not to aridity.

? The geographical pattern of genetic variation found in morphology and biomass allocation did not translate into population differences in drought tolerance or phenotypic plasticity to water availability, indicating that water availability is not a likely driver of the regional variation observed in the studied traits of P. pinaster at the seedling stage.

     

The combined effects of thinning and prescribed fire on carbon and nutrient budgets in a Jeffrey pine forest

? Both burning and harvesting cause carbon and nutrient removals from forest ecosystems, but few studies have addressed the combination of these effects. For a Pinus jeffreyii forest in the Sierra Nevada Mountains of California, we posed the question: what are the relative impacts of thinning and subsequent burning on carbon and nutrient removals?

? The thinning methods included whole-tree thinning (WT, where all aboveground biomass was removed) cut to length (CTL, where branches and foliage were left on site in a slash mat on top of skid trails) and no harvest (CONT). Total C and nutrient exports with thinning and burning were greater in the WT and CTL than in the CONT treatments. Total C and N removals were approximately equal for the WT and CTL treatments, although harvesting dominated exports in the WT treatment and burning dominated exports in the CTL treatment. Total removals of P, K, Ca, Mg and S were greatest in the WT treatments, where harvesting dominated removals.

? Comparisons of nutrient removals with ecosystem capital and calculations of potential replenishment by atmospheric deposition suggested that N is the nutrient likely to be most depleted by harvesting and burning treatments.

     

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.

     

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.

     

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.

     

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.

     

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.

     

Forest floor contribution to phosphorus nutrition: experimental data

? Although accumulation of decomposing litter temporarily removes nutrients from active circulation, it creates a medium that is more suitable for nutrient uptake where soil conditions are unfavorable.

? A pot experiment was conducted using labeling of isotopically exchangeable phosphate ions of the soil and applying the dilution principle to accurately assess the contribution of the forest floor to P nutrition of maritime pine seedlings (Pinus pinaster Aït.). Three-week-old maritime pine seedlings were planted in pots containing either mineral soil (MS) or mineral soil covered with a forest floor layer (MS+FF).

? After 130 d, P uptake was still insignificant in the MS treatment while the P content of the seedlings in the MS+FF treatment increased tenfold with respect to the initial P content. In the latter treatment, the forest floor contributed 99.1% of the P supply to pine seedlings.

? The higher P uptake from the forest floor than from the mineral soil may be explained by its lower ability to retain inorganic P, which enabled a higher concentration of inorganic P to be maintained in solution.

     

Genetic variation, mating patterns and gene flow in a Pinus pinaster Aiton clonal seed orchard

? Relatedness among parents, variation in clonal fertility and background pollination deviate the realized genetic gain and the gene diversity of open pollinated seed orchard from expectation, in particular in wind pollinated species such as Pinus pinaster Aiton.

? This work investigates the genetic variation, the mating system and the pollen contamination in a P. pinaster clonal seed orchard (CSO), by screening the 60 clones from the CSO and the seeds collected from 21 mother-trees with three nuclear microsatellites.

? The expected diversity was similar, but the observed heterozygosity decreased 20% in the progenies compared with the parental trees. The outcrossing rate was 90.1%, the biparental inbreeding 21.7% computed through a multilocus approach, and the observed selfing 3.9%. The observed gene flow from outside the CSO was 52.4%.

? From the results we concluded that the observed gene flow and the biparental inbreeding were high, and care should be taken in the implementation and management of future CSO, in particular clones should be checked for relatedness and the ramet number could be directly proportional to their breeding value.

     

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.

     

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.

     

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.

     

Circumferential variation of heartwood and stem quality in maritime pine stems

Maritime pine (Pinus pinaster Ait.) is an important timber and pulpwood species covering about 4 million ha in Portugal, Spain and France. This work studied the cross-sectional distribution of heartwood and sapwood along the stem using 3D modelling on 20 trees. The target was to assess the potential of this species for the industrial production of heartwood sawn products. The maritime pine stems were characterized by wide sapwood of 10 cm, in average, and the presence of heartwood at all height levels with constant diameter up to 35 % tree height, decreasing afterwards. In the radial section, the heartwood follows a circumferential-like shape which changes along the stem. Pith eccentricity index values were low and homogeneous regarding to stem and higher for heartwood. Heartwood cross-sectional shape was constant up to 45 % of total height of the tree and more elongated upwards. The constant heartwood diameter, low taper and pith eccentricity index, and an approximate circular shape up to 10 m height indicated these species’ potential to produce solid wood products made out of heartwood.     

Manipulating nutrient and water availability in a maritime pine plantation: effects on growth, production, and biomass allocation at canopy closure

? We present here the results of a water and nutrient manipulation experiment in a five-year-old plantation of maritime pine in south-western France.

? Water and nutrient levels were manipulated in a factorial design with two levels of irrigation (control receiving only rainfall (C) and irrigated (I)) and three levels of fertilisation (control with no added nutrients (C), P-only (P) and annual addition of a complete nutrient mix (F)) in order to quantify growth limitations of plantation forest in this particular area.

? The treatments applied during five years increased aboveground biomass annual increment by 4% (I) to 58% (IF) with respect to the control (C). The fertilised plots had a slightly non-significant lower root-to-shoot ratio. The effect of irrigation was maximal in 2002, resulting in 6%, 7% and 12% higher growth rate on the F, C and P plots, respectively. A windstorm disturbed the experiment in 1999 and has affected preferentially the fertilised plots, with IF plots displaying 60% damage.

? The higher growth rate of fertilised and irrigated plots was attributed to both an increase (estimated at 5 to 15%) in the amount of light absorbed by the canopy, and an increase (estimated at 26% for IF plots) in the amount of above-ground biomass produced annually per unit leaf area.