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.

     

Influence of several tree traits on rainfall partitioning in temperate and boreal forests: a review

? In forests, rainfall is partitioned into intercepted water (IW), throughfall (TF) and stemflow (SF). We reviewed the majority of published works comparing water flows among tree species in temperate and boreal forests to test the effect of several tree traits on water flows.

? We hypothesized that water flows differed between evergreen and deciduous species, and according to successional status and bark roughness. We also investigated that water flows would be explained by stand variables such as basal area.

? Linear mixed models fitted on reviewed data showed that evergreens had a lower TF than deciduous trees (decrease of 13.9% of total precipitation year-round and 8.4% over the growing period). Similar results were found for conifers compared to broadleaves. TF also declined with the successional status from pioneer to late-successional tree species. SF decreased with bark roughness from smoother to rougher bark. Evergreens had water flows that were dependent on age of the stand, especially for TF which increased by 15.6% of total precipitation from young to adult forests.

? The large scale of TF differences between tree genera together with specific transpiration amounts and rooting features highlighted in other studies should result in significant differences in soil water content among tree species. This may have consequences on stand fitness and growth, and understory vegetation.

     

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.

     

Evaluation of decay resistance of wood products made from borax-impregnated wood and bonded with a formaldehyde-free cornstarch and tannin adhesive

? At present, the production of wood composites mainly relies on the petrochemical-based and formaldehyde-based adhesives such as phenol-formaldehyde (PF) resins and urea-formaldehyde (UF) resins, which are non-renewable and therefore ultimately limited in supply.

? This paper concerns the decay resistance of wood products bonded with a new, environment-friendly adhesive derived from abundant and renewable cornstarch and tannin. To improve the total resistance of the composite against both Coriolus versicolor and Coniophora puteana rot fungi, borax (di-sodium tetraborate) was added in proportions of 0.5%, 1% and 2% (w/w) to the cornstarch-tannin adhesives.

? The results show that increasing the concentration of borax in the adhesive decreased the mechanical properties of the composite. The best way to avoid this problem was to use wood impregnated with borax.

? Biodegradation studies were conducted on new composites, first without any treatment, followed by borax at 0.5% aqueous solution treatment. The results show that wood impregnated with borax, in the presence of tannin and sodium hydroxide in the adhesive improves the total resistance of the wood composite against both Coriolus versicolor and Coniophora puteana rot fungi.

     

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.

     

Frost damage in Pinus sylvestris L. stems assessed by chlorophyll fluorescence in cortical bark chlorenchyma

? Several techniques were used to investigate the threshold minimum temperature inducing severe frost damage during winter in Scots pine (Pinus sylvestris L.) needles and stems from southern provenances in the Iberian Peninsula.

? Chlorophyll (Chl) fluorescence, electrolyte leakage (EL), visual scoring (VS), and the normalized difference vegetation index (NDVI) were compared.

? Chl fluorescence, and in particular the maximum potential photosystem II (PSII) efficiency — estimated through the dark-acclimated F_V/F_M ratio-, was found to be a simple, non-destructive indicator of freezing tolerance in needles, as expected.

? Moreover, the existence of a Chl-containing tissue, the cortical bark chlorenchyma, allowed us the use of Chl fluorescence as a new non-destructive indicator of frost damage in live tissues of Pinus sylvestris L. stems.

? Freezing tolerance values of southern provenances are compared with those found by other authors in the northern distribution limit of the species.

     

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.

     

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.

     

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.

     

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.

     

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 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).

     

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.

     

Variations of plant and soil 87Sr/86Sr along the slope of a tropical inselberg

? From the summit downslope a granitic inselberg in French Guiana, soils and vegetation evolve from bare granite covered by cyanobacteria, to a savannah-type vegetation on thin patchy sandy accumulations, then to a low forest on shallow young soils and to a high forest on deep highly weathered ultisols.

? We have used element budgets and Sr isotopic variations in soils and plants to investigate the mineral nutrient supply sources of the different plant communities.

? Granite and atmospheric deposition have ⁸⁷Sr/⁸⁶Sr ratios of 1.3 and 0.71, respectively. The ⁸⁷Sr/⁸⁶Sr ratio of cyanobacteria (0.72) suggests granite weathering by cyanobacteria crusts. The ⁸⁷Sr/⁸⁶Sr ratio of the savannah-type vegetation is 0.73 and varies between 0.75 and 0.76 in the low and high forest leaf litter regardless of soil depth, age and degree of impoverishment.

? These almost constant ratio suggest that forest Sr comes from rainwater and from the summit of this inselberg, where it is released and redistributed along the slope, by surface flow, lateral redistribution of litter, and mineral particles. However, because of its very low content in the rock and soils, Ca is supplied to plants by atmospheric deposition.

     

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.

     

Comparing floristic composition in treefall gaps of certified, conventionally managed and natural forests of northern Honduras

? Forest certification sets requirements for minimizing the impacts of logging on the natural structure and floristic composition of forests.

? We assessed the impact of certification by comparing the floristic composition of 52 taxa of trees and shrubs in the treefall gaps of certified, conventionally managed and protected forests in northern Honduras.

? The highest abundance of light-benefiting taxa was found in certified forests, whereas conventionally managed forests were floristically more similar to natural forests. The environmental conditions measured in certified gaps were not favourable for a natural forest floristic composition.

? Past logging may have altered the species composition in certified forests relatively more than in conventionally managed forests. This implies that the need for restoration operations should be considered in certification requirements, along with landscape-level planning to enhance post-logging recovery.

     

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.

     

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.

     

Winter variation in physiological status of cold stored and freshly lifted semi-evergreen Quercus nigra seedlings

? Water oak (Quercus nigra L.) is a tardily deciduous species commonly planted in afforestation projects in the Lower Mississippi River Alluvial Valley, USA. Field performance is often marked by low survival rates and top dieback, which may be associated with poor physiological quality of planting stock.

? We investigated physiological status of cold stored (2–4 °C; CS) and freshly lifted (FL) seedlings during the period between lifting and planting (December — February). In mid-February, seedlings were transplanted into a controlled greenhouse environment for 90 d to evaluate post-transplant growth performance.

? Net photosynthetic rates were positive until late January (generally greater in CS seedlings) and became negative thereafter. FL seedlings generally had lower LT₅₀ values from freeze-induced electrolyte leakage (FIEL), reflecting greater cold hardiness. FIEL of foliage provided the best indicator of physiological status, though terminal buds may serve as a suitable substitute. All seedlings experienced top dieback following transplant; CS seedlings had less relative root-collar diameter, height, and root volume increments.

? Cold storing seedlings did not appear to prolong dormancy, increase stress resistance, or hold promise as a means to improve outplanting success. Regardless of storage regime, seedlings appeared to be most cold hardy and perhaps stress resistant until late January.

     

X-ray tomography as a tool for detailed anatomical analysis

? Wood identification, anatomical examination and retrieval of quantitative information are important aspects of many research disciplines. Conventional light microscopy with a camera and (semi)-automatic image analysis software is an often used methodology for these purposes. More advanced techniques such as fluorescence, scanning electron, transmission electron, confocal laser scanning and atomic force microscopy are also part of the toolset answering to the need for detailed imaging.

? Fast, non-destructive visualization in three dimensions with high resolution combined with a broad field of view is sought-after, especially in combination with flexible software.

? A highly advanced supplement to the existing techniques, namely X-ray sub-micron tomography, meets these requirements. It enables the researcher to visualize the material with a voxel size approaching < 1 μm for small samples (< 1 mm). Furthermore, with tailor-made processing software quantitative data about the wood in two and three dimensions can be obtained. Examples of visualization and analysis of four wood species are given in this paper, focusing on the opportunities of tomography at micron and sub-micron resolution.

? X-ray computed tomography offers many possibilities for material research in general and wood science in specific, as a qualitative as well as a quantitative technique.