Vessel size and number are contributors to define wood density in cork oak

Cork oak (Quercus suber L.) has a dense wood that allows high-quality uses. In the present work, we study the influence of vessel characteristics, measured through image analysis and optical microscopy, on wood density, measured using X-ray microdensitometry, on 40-year-old trees. Vessel area increases with cambial age (5403–33064 μm²), while wood density decreases (1.229–0.836 g/cm³). The number of vessels is relatively constant at 6 vessels/mm², while vessel proportion in cross-section increases from 3.3% near the pith to 20.5% near the bark. In growth rings closest to the pith, with high wood density and low vessel area, the relationship between the two variables is linear (R ² = −32.1%, P < 0.01) but with increasing tree age and vessel size, the wood density remains rather constant, suggesting that decreases in density might compromise mechanical support of the tree at a stage when the increase in cross-sectional area alone might not provide mechanical stability. Other anatomical characteristics not considered in this study, like large xylem rays that increase with cambial age, may be responsible for the constant density.     

Clues to wood quality and production from analyzing ring width and density variabilities of fertilized Pinus taeda trees

Tree growth and wood density are influenced by forest management. Nevertheless, few studies have evaluated their variability responses to fertilizer treatments at inter-annual, inter-tree and stand-production levels. Therefore, the annual ring width (RW) and density (RD) of sixty 17-year-old-Pinus taeda trees fertilized with six doses of composted pulp-mill sludge (CPMS) were analyzed. Ten trees for each treatment were felled and from which wood discs were taken at different trunk heights. The annual RW and RD were provided by X-ray microdensitometry, synchronized and the trunk basic specific gravity (SG_B) and biomass calculated. The effects of CPMS treatments were explored using interaction of variables RW and RD with cambial age, diameter, trunk SG_B and biomass production. Trees treated with CPMS grow faster, increasing their biomass (up to 108%), presenting lower wood density values (significant up to the 6th year) and reaching the mature wood later than untreated trees. Furthermore, the potential use of RW and RD in allometric equations showed good accuracy to predict trunk SG_B and biomass. Altogether, our results indicated that ring width and density revealed the impacts of fertilization treatment on wood quality and production. Our study also provides useful information for forest managers on the fertilization monitoring process.

     

Inter-tree and intra-tree variations in ring width and wood density components in balsam fir (Abies balsamea)

This study quantified and compared intra-tree and inter-tree variations in ring width and wood density components in balsam fir (Abies balsamea) grown in Quebec, Canada. In addition, the study examined correlations between ring width and wood density components at different stem positions from the stump level to the stem top. Ring width and wood density components of individual rings were measured by X-ray densitometry. Both the intra- and inter-tree variations in balsam fir are large, but the inter-tree variation is relatively smaller than the intra-tree variation. Much of the intra-tree variation is due to the radial variation, whereas the axial variation is much smaller. Compared to ring width and its components, wood density characteristics show a considerably smaller variation at both the inter- and intra-tree level. In almost all wood characteristics studied (except for latewood width), the intra- and inter-tree variations are more or less influenced by tree age. Cambial age explains more intra-tree variation in wood density components than ring width, whereas more intra-tree variation in ring width components is due to ring width. Cambial age and ring width explain a comparable percentage of variation in ring density. Only a few of the correlations between ring width and wood density components vary significantly with stem position from the stump to the stem top at the inter-tree level. In balsam fir, a negative correlation between ring density and ring width is significant in the butt log, but the correlation decreases to an insignificant level at and above a height of 3.0 m.     

Variations and correlations of various ring width and ring density features in European oak: implications in dendroclimatology

Summary Variations and correlations of various ring width and ring density features were analyzed in 18 European oak (Quercus petraea andQuercus robur) trees from northeastern France. In light of these analyses, the goodness of various tree-ring features as a climatic parameter was discussed. In general, ring density features (viz. earlywood density, latewood density, average ring density, minimum density and maximum density) show a stronger response to calendar year, a comprehensive climatic variable, than ring width features (viz. earlywood width, latewood width, total ring width and latewood percent). The response of latewood features (viz. latewood width, latewood density and maximum latewood density) is stronger than that of earlywood features (viz. earlywood width, earlywood density and minimum earlywood density). Average ring density seems to be the most sensible tree-ring feature in European oak in terms of the response to calendar year. Moreover, total ring width as a climatic parameter is not as good as latewood width, and maximum (latewood) density and minimum (earlywood) density appears not to contain as much climatic information as (average) latewood density and (average) earlywood density, respectively.This project was supported by the institut National de la Recherche Agromonique (INRA). Thanks are due to Dr. G. Nepveu for his constant encouragement and assistance during this study     

Within and between-tree variation in the biometry of wood rays and fibres in cork oak (Quercus suber_L.)

The variability of fibre and ray dimensions was studied at three radial positions (10, 50 and 90% of radius) in 5 cork oaks (Quercus suber L.) approximately 40 years old. Uniseriate ray height and cell number as well as multiseriate ray height and width were measured in tangential sections. Fibre length, width and wall thickness were measured after maceration. The most striking features of cork oak wood were the large multiseriate rays and the frequent homocellular uniseriate rays. On average multiseriate rays were 5.16 mm high and 0.48 mm wide, and uniseriate rays contained 11 cells and were 227 μm high. Neighbour rays presented large dimensional differences but there was no radial variation and there were very small between-tree differences. The radial maintenance of high and wide rays was discussed as a means to improve water shortage in response to water stress. Fibre dimensions increased radially but differed little between trees: on average, 960–1,220 μm length, 18.40–21.49 μm width and 6.66–8.07 μm wall thickness. The ray pattern showed enough variation for visual appreciation while ensuring within and between-tree uniformity. The presence of rays has to be taken into consideration for designing wood drying programs.     

The effect of cork removal on the radial growth and phenology of young cork oak trees

The effect of the removal of cork was studied in 11-years old cork oak trees (Quercus suber L.), growing in favorable conditions, in relation to phenology and radial growth during two years. Longevity of leaves was 14–15 months (1996, 1998) and 10 months (1997). Bud burst started in mid-February and leaf flushing in April, extending until June. Neither a distinct two-period flushing nor an autumn bud burst or leaf flushing were observed. Radial growth started in mid-April and continued until the end of November, with the maximum growth in June and July. In trees with the removal of cork, leaf abscission occurred a little earlier and new branches had on average 23% fewer new leaves. The radial growth of the trees and its general seasonal pattern were not affected by cork removal in the year of removal or in the year after. The only observation was a shift of the maximum radial growth rate from June to July for the trees where cork had been removed.     

Variation in wood density and ring width in Acacia melanoxylon at four sites in Portugal

The radial variation in wood density in Acacia melanoxylon R. Br. was studied using microdensitometry by sampling 20 trees with a 40-cm diameter class at four sites in Portugal. The measurements were taken from pith to bark at breast height. A. melanoxylon had an average ring density of 0.607 g cm^?3, ranging from 0.556 to 0.630 g cm^?3. The mean growth was 6.0 mm year^?1. Latewood corresponded, on average, to 34 % of the ring width. Between-tree variability at each site was the main source of variation in the density components, representing between 30 and 54 % of the total variation. Between-sites variability represented from 0 to 21 % of the total variation in density components. The environmental effects (site related) were more pronounced on latewood, while the genetic effect (tree related) was more evident in earlywood. Ring width, latewood percentage and heterogeneity index were independent from site, trees in site and age effects. The values of wood density and radial growth revealed that A. melanoxylon can be important as a commercial timber species in Portugal.     

Influence of cambial age and climate on ring width and wood density in Pinus radiata families

? Context

The correlation between tree ring width and density and short-term climate fluctuations may be a useful tool for predicting response of wood formation process to long-term climate change.

? Aims

This study examined these correlations for different radiata pine genotypes and aimed at detecting potential genotype by climate interactions.

? Methods

Four data sets comprising ring width and density of half- and full-sib radiata pine families were used. Correlations with climate variables were examined, after the extraction of the effect of cambial age.

? Results

Cambial age explained the highest proportion of the ring to ring variation in all variables. Calendar year and year by family interaction explained a smaller but significant proportion of the variation. Rainfall had a positive correlation with ring width and, depending on test site, either a negative or positive correlation with ring density. Correlations between temperature during growing season and ring density were generally negative.

? Conclusion

Climate variables that influence ring width and wood density can be identified from ring profiles, after removing the cambial age effect. Families can be selected that consistently show desirable response to climate features expected to become prevalent as a result of climate change.     

Differences of raw cork quality in productive cork oak woods in Sicily in relation to stand density

Differences of raw cork quality, in terms of thickness and porosity, were observed in productive cork oak (Quercus suber L.) stands in the Nebrodi and Iblei mountains of Sicily in relation to stand density. Five study areas were chosen across a range of different stand density satisfying specific sampling requirements among two managed cork oak woods.Analysis of variance using Fisher–Snedecor‘s F test (p < 0.05) were used to identify statistically significant differences of cork quality and sylvicultural, dendrometric and cork stripping parameters between study areas within each stand. Relationships between cork quality and sylvicultural and dendrometric parameters were also tested using Pearson's correlation coefficient (r). Results shown that sylvicultural practices are a key factor influencing cork quality: statistically significant differences in cork thickness and crown area were observed between areas at different stand density; positive correlations were found between cork thickness and crown area in lower density study areas favouring a significant vegetative activity; negative correlations were found between parameters of cork stripping (cork stripping coefficient, cork stripping surface, maximum height of cork stripping) and cork thickness. No correlations were found between cork porosity and dendrometric or sylvicultural parameters, demonstrating the probable genetic origin of cork porosity. Results shown that cork quality parameters and stand attributes may be used in sustainable management models of cork oak forests.     

Radial variation of wood properties in Neolamarckia cadamba trees from an East Java community forest

Radial variations in wood properties, cell morphologies and cell proportions were investigated for nine 4-year-old Neolamarckia cadamba trees planted in a community forest in East Java, Indonesia. The relationships between stem diameter and wood properties, cell morphologies or cell proportions were determined to ascertain the effects of stem diameter on these properties. In the radial variations, almost all of the wood properties, cell morphol- ogies and cell proportion in N. cadamba were changed at around 4 to 6 cm distance from the pith, except for fibre diameter, and the proportion of fibre and axial parenchyma. For trees with larger stem diameter, the vessel diameter was also found to increase. In addition, the ray parenchyma and cell wall proportions in trees with larger stem diameter were lower than those in trees with smaller stem diameter. However, the wood properties obtained from the larger stem diameter trees did not always show lower values. The wood properties in N. cadamba were correlated to fibre wall thickness and vessel diameter.     

Radial variation of vessel size and distribution in cork oak wood (<Emphasis Type="Italic">Quercus suber</Emphasis> L.)

Quercus suber L. is an important species producing cork whose wood characteristics have not been investigated a lot. Cork oak wood vessels are a striking feature and the most abundant wood tissue largely influencing density and permeability. Vessel size and distribution were studied in approximately 40 year-old and never debarked cork oaks by continuously measuring along the radial direction in the transverse section of wood discs taken at 1.3 m of height using image analysis techniques. The vessel size increases with age from 7660 ± 2286 to 21136 ± 6119 μm², the conductive area from 5.4 ± 2.2 to 11.6 ± 3.9%, and the vessel density remains approximately constant between 5.2 ± 1.5 and 7.3 ± 3.5 vessels/mm². In comparison with ring-porous and some evergreen oaks, cork oaks show a similar conductive area but smaller vessels. Vessel architecture is known to play an important role on oaks tolerance to hydric stress, and these cork oak trees were growing under very harsh edaphoclimatic conditions, not tolerated by other oaks. The well-developed and deep root system allowing access to constant water supply may contribute to the cork oak’s relatively high conductive area.     

Stem modeling and simulation of conversion of cork oak stems for quality wood products

Recent research is underway to study cork oak (Quercus suber L.) wood potential for production of wood components. A total of 35 stems of young and mature cork oaks were sampled, live sawn into flitches, scanned using VTT’s WoodCIM^®, and the measured data computed by VTT’s PuuPilot software, allowing stem 3D reconstruction. Sawing simulations were run for 0.5 m and 1 m logs and the whole stem. Sawn products were (1) planks, (2) parquet, (3) lamparquet, and (4) external component of multilayer planks. Cork oak stems showed a moderate to small taper (mean 24 mm/m). Curviness varied between straight to significantly crooked stems (mean value 40 mm) Batch yields for the tested products ranged 25–43% and 37–50% for 0.5 m logs of young and mature trees, respectively; for 1 m logs, batch yields ranged 19–41% and 25–54%. When using the whole stem, batch yields were lower, ranging 11–38% and 15–50%. Higher yields were obtained for all log lengths and samples for production of lamparquet, parquet, and multilayer component.     

Factors affecting post-fire crown regeneration in cork oak (Quercus suber L.) trees

Cork oak (Quercus suber) forests are acknowledged for their biodiversity and economic (mainly cork production) values. Wildfires are one of the main threats contributing to cork oak decline in the Mediterranean Basin, and one major question that managers face after fire in cork oak stands is whether the burned trees should be coppiced or not. This decision can be based on the degree of expected crown regeneration assessed immediately after fire. In this study we carried out a post-fire assessment of the degree of crown recovery in 858 trees being exploited for cork production in southern Portugal, 1.5 years after a wildfire. Using logistic regression, we modelled good or poor crown recovery probability as a function of tree and stand variables. The main variables influencing the likelihood of good or poor crown regeneration were bark thickness, charring height, aspect and tree diameter. We also developed management models, including simpler but easier to measure variables, which had a lower predictive power but can be used to help managers to identify, immediately after fire, trees that will likely show good crown regeneration, and trees that will likely die or show poor regeneration (and thus, potential candidates for trunk coppicing).     

Modelling cork oak production in Portugal

The layout of 32 permanent plots in the most important cork production regions of Portugal and the measurement of the most representative tree parameters were performed for statistically sound cork weight modelling. The establishment of cork weight equations is presented for the different cork oak study areas considered alone or grouped according to the most representative tree parameters. For the presented cork weight models was chosen independent variable the total stripping surface (artmax). For the selection criteria five statistic were used, including PRESS, APRESS and Mallows's statistic. The crossvalidation of the best classified models was performed and to discuss the importance of the “study area” factor, a regression analysis with “dummy” variables was performed.     

Physiological responses of cork oak and holm oak to infection by fungal pathogens involved in oak decline

The aim of this research was to study the changes in net photosynthesis and stomatal conductance values in 3‐year‐old cork oak and holm oak seedlings growing in natural conditions and inoculated with Apiognomonia quercina, Biscogniauxia mediterranea, Botryosphaeria corticola and Pleurophoma cava. Throughout the 4‐month experimental period, the evolution of visual external symptoms and the values of physiological variables were periodically recorded. All pathogens caused stem lesions around the infection point; however, the lesions caused by B. corticola were longer in both oak species. On cork oak seedlings, all pathogens induced a significant and gradual reduction in net photosynthesis and stomatal conductance values, whereas other physiological disturbances were induced only by B. corticola infections on holm oak seedlings.     

Variables influencing cork thickness in Spanish cork oak forests: A modelling approach

In this study, we evaluate the influence of different variables on cork thickness in cork oak forests. For this purpose, first we fitted a multilevel linear mixed model for predicting average cork thickness, and then identified the explanatory covariates by studying their possible correlation with random effects. The model for predicting average cork thickness is described as a stochastic process, where a fixed, deterministic model, explains the mean value, while unexplained residual variability is described and modelled by including random parameters acting at plot, tree, plot × cork harvest and residual within-tree levels, considering the spatial covariance structure between trees within the same plot. Calibration is carried out by using the best linear unbiased predictor (BLUP) theory. Different alternatives were tested to determine the optimum subsample size which was found to be appropriate at four trees. Finally, the model was applied and its performance in the estimation of cork production was tested and compared with the cork weight model traditionally used in Spain.     

Fingerprinting and genetic variability in cork oak (Quercus suber L.) elite trees using ISSR and SSR markers

Quercus suber L., is a socially, economically and ecologically important forest species in rural areas of the Mediterranean basin. Fifty three elite-trees from nine stands of four provenance regions in the Community of Extremadura (Spain) were analysed with the aim to establish their DNA-fingerprinting and the genetic relationships among them. Two types of molecular markers, microsatellites and intermicrosatellites, were used for tree genotyping. Microsatellite markers could distinguish 94% of the trees. Observed and expected heterozygosities, and effective number of alleles were correlated with the polymorphism information content (PIC). Intermicrosatellite patterns were used to construct a dendrogram. They showed high levels of genetic diversity in these elite trees, without any clear relationship with provenance regions. Both types of markers detected a high level of intra-stand variability, which is common in Quercus species, probably due to its mating system, the low distance among stands and the small size of stands. Others factors that could affect this result, such as introgression between Q. suber L. and Q. ilex L., are also commented. The high level of genetic variability detected in these elite trees can be useful for improvement programs. Also the usefulness of SSR and ISSR markers to establish the DNA-fingerprinting of each tree could be focused to select clonal lines of commercial interest.     

Analysis of spatial patterns of oak decline in cork oak woodlands in Mediterranean conditions

? Cork oak mortality is a recurrent problem in southwestern Portugal. Despite the perception of increasingly visible damage in oak woodlands on drought-prone sites, the role of the various environmental factors in their decline is not clear.

? To describe the spatial patterns of cork oak (Quercus suber L.) mortality, a cork oak mortality index (MI) was determined for each landscape feature (agroforestry system, soil type, slope and aspect) using a GIS approach. To achieve this goal, a logistic regression model was formulated analyzing interactions between landscape attributes and allowing a prediction of cork oak mortality.

? Maximum values of MI were found in (i) shrublands and open woodlands with shrub encroachment (MI 6 and 3, respectively), where competition for soil water between tree and understory increases; and (ii) on lower slopes in the rounded hilltops and smooth hillsides or shallow soils where access to groundwater resources during summer drought is difficult.

? The model highlighted the importance of the agroforestry systems on cork oak mortality and may be used to identify sensitive areas where mitigation actions should be employed in a scenario of increasing drought severity in these Mediterranean ecosystems.