Philippe Rozenberg

Genetic parameters of growth and wood quality traits in Picea abies

Genetic parameters were estimated for wood and growth traits in two 19-yr-old clonal trials and a 40-yr-old full-sib progeny trial of Norway spruce [Picea abies (L.) Karst.]. In the clonal trials high (>0.4) broad-sense heritabilities were found for wood density traits, lignin content, number of internal cracks, growth traits, spiral grain and number of resin canals. Moderate (0.2–0.4) heritabilities were found for tracheid lumen diameter and cell wall thickness, microfibril angle and tracheid length, while low heritabilities (<0.2) were found for pulp yield, fibre strength, wood stiffness and wood colour. Lignin content and pulp yield showed low genetic variation, whereas the genotypic coefficient of variation for most other traits ranged between 5 and 15%. Most traits showed low levels of genotype by environment interaction. Among the wood properties, latewood proportion, earlywood density and ring density showed significant, adverse correlations with volume in both clonal trials.

Colour of larch heartwood and relationships to extractives and brown-rot decay resistance

Larch heartwood is appreciated for its good mechanical properties, its colour and its texture, and it is often used outdoors because of its natural durability (decay resistance). In this study the colour of larch heartwood was studied in relation to extractives and decay resistance, with the aim to estimate durability of larch heartwood from its colour. On a total of 293 trees colour in the CIE L*a*b* space (L* lightness, a* red/green axis, b* yellow/blue axis), extractives content (acetone and hot-water extractives, amount of phenolics) and the brown-rot decay resistance were determined. For calculating the relative decay resistance ( x), mass loss after inoculation for 16 weeks with two fungi [ Coniophora puteana (Schum.ex.Fr.) Karst., Poria placenta (Fr.) Cke, European standard EN 113] of larch heartwood samples was compared to Scots pine ( Pinus sylvestris L) sapwood reference samples (EN 350-1). Different species [Japanese larch ( Larix kaempferi Lamb.), Hybrid larch (Larix deciduax L. kaempferi) and European larch ( L. decidua Mill.)], provenances and age classes (38-year, >150-year) were included. Japanese larch heartwood turned out to be significantly more reddish (higher a*-values) compared to the European larch provenances. Reddishness of the hybrids was intermediate. The red hue (+a*) was strongly correlated with the amount of phenols ( r =0.84) and decay resistance ( r =0.63) and therefore suitable for prediction of both parameters. The results suggest that colour measurements of larch heartwood could be of benefit in tree breeding programs and for an optimised utilization of larch timber.

QTLs and candidate genes for wood properties in maritime pine (Pinus pinaster Ait.)

A three-generation outbred pedigree of 186 individuals was used to identify the genomic regions involved in the variability of chemical and physical wood properties of Pinus pinaster. A total of 54 quantitative trait loci (QTLs) was detected, with an average of 2.4 QTLs per trait. Clusters of wood properties QTLs were found at several points in the genome, suggesting the existence of pleiotropic effects of a limited number of genes. The co-localizations observed in this study are in accordance with the genetic correlations previously reported in the literature. In addition, in an attempt to identify the genes underlying the QTLs, nine wood quality candidate genes involved in cell wall structure were localized on the genetic map. Only one of them, Korrigan, a gene encoding for a β 1-4 endo-glucanase known in Arabidopis thaliana to be involved in polysaccharide biosynthesis, co-localized with a wood quality QTL cluster involved in hemicellulose content and fibre characteristics. This finding is in accordance with results previously reported for this gene regarding its expression variability (transcriptome and proteome levels) and patterns of molecular evolution. The pertinence of this result will be tested in more rigorous designs in order to identify early selection predictors for wood quality.

Comprehensive genetic dissection of wood properties in a widely-grown tropical tree: Eucalyptus

BackgroundEucalyptus is an important genus in industrial plantations throughout the world and is grown for use as timber, pulp, paper and charcoal. Several breeding programmes have been launched worldwide to concomitantly improve growth performance and wood properties (WPs). In this study, an interspecific cross between Eucalyptus urophylla and E. grandis was used to identify major genomic regions (Quantitative Trait Loci, QTL) controlling the variability of WPs.ResultsLinkage maps were generated for both parent species. A total of 117 QTLs were detected for a series of wood and end-use related traits, including chemical, technological, physical, mechanical and anatomical properties. The QTLs were mainly clustered into five linkage groups. In terms of distribution of QTL effects, our result agrees with the typical L-shape reported in most QTL studies, i.e. most WP QTLs had limited effects and only a few (13) had major effects (phenotypic variance explained > 15%). The co-locations of QTLs for different WPs as well as QTLs and candidate genes are discussed in terms of phenotypic correlations between traits, and of the function of the candidate genes. The major wood property QTL harbours a gene encoding a Cinnamoyl CoA reductase (CCR), a structural enzyme of the monolignol-specific biosynthesis pathway.ConclusionsGiven the number of traits analysed, this study provides a comprehensive understanding of the genetic architecture of wood properties in this Eucalyptus full-sib pedigree. At the dawn of Eucalyptus genome sequence, it will provide a framework to identify the nature of genes underlying these important quantitative traits.

Can wood density be efficiently selected at early stage in maritime pine (Pinus pinaster Ait.)

Growth and stem straightness are the two main selection criteria in the French maritime pine breeding programme. In this contribution, our objective was to study the possibility of selecting for wood density assessed at an early stage. We measured X-ray wood density, in three progeny tests, on more than 1 900 breast high increment cores. High relative expected genetic gain (from 3 to 9%) was found for wood density at mature stage. Juvenile and adult wood density estimations were well-correlated; the relative efficiency of early selection reached about 80% at 12 years old. As it would not be realistic to use the X-ray densitometer at an operational scale in a breeding programme, we tested the efficiency of using the Resistograph, an indirect method to assess wood density on live trees. High correlation between the Resistograph and density data (R2 = 0.93 on familial data) was found, suggesting that wood density through Resistograph assessment could be soon integrated as a new selection criterion in our breeding programme.RésuméLa croissance et la rectitude du tronc sont les deux principaux critères de sélection dans le programme d’amélioration génétique du pin maritime en France. Cette étude a pour objectif d’évaluer la possibilité de sélectionner la densité du bois au stade juvénile. Plus de 1 900 arbres, provenant de trois tests, ont été analysés par microdensitométrie. Des gains génétiques relatifs élevés ont été mis en évidence pour la densité du bois au stade adulte (3 à 9 %). Les estimations de la densité réalisées aux stades adulte et juvénile étant bien corrélées, l’efficacité relative de la sélection atteint environ 80 % à l’âge de 12 ans. Les mesures de densité par rayons X ne sont pas envisageables à grande échelle dans le cadre d’un programme d’amélioration, ainsi nous avons évalué l’efficacité de l’utilisation du Résistographe, une méthode indirecte pour estimer la densité du bois sur arbres debout. Des corrélations élevées ont été mises en évidence entre les données obtenues avec le Résistographe et celles obtenues par microdensitométrie (R2 = 0.93 avec les données familiales). Il semble donc possible d’intégrer la densité du bois, estimée avec le Résistographe, comme nouveau critère de sélection dans notre programme d’amélioration.

Genetic variation of xylem hydraulic properties shows that wood density is involved in adaptation to drought in Douglas-fir (Pseudotsuga menziesii (Mirb.))

IntroductionRelationships between wood density and hydraulic efficiency and safety (hydraulic specific conductivity and vulnerability to cavitation, respectively) could clarify the physiological process explaining the impact of density on fitness. We have used new, relatively high-throughput phenotyping methods to estimate genetic variation of wood hydraulic specific conductivity (ks) and vulnerability to cavitation (VC) as an important step toward demonstrating the adaptive value of wood density.ObjectiveThe first aim of this study is to test if, in Douglas-fir, there is a relationship between wood hydraulic properties (ks and VC) and wood density. The second objective is to estimate genetic variation of wood ks and VC. These results could aid understanding of the role of wood density in the hydraulic properties of xylem and may clarify the role of wood density in adaptation to drought.ResultsMany significant relationships were found between wood density and wood hydraulic properties at clone and tree level, as well as significant genetic variation for ks and VC. We have also found positive correlations between tree height, specific conductivity and vulnerability to cavitation, but no relation was found between radial growth and hydraulic variables.ConclusionsOur results suggest that wood density has an adaptive value and that microdensity can be used to study adaptation to drought in Douglas-fir. The novel methods used to measure ks and VC proved to be interesting alternatives for localized measurement of wood hydraulic properties and were compatible with a robust estimation of genetic variation.

Improvement of Pinus pinaster Ait elite trees selection by combining near infrared spectroscopy and genetic tools

Abstract The first and the second generation of improved maritime pine (Pinus pinaster) have produced gains of up to 30% in stem volume, a reduction of final harvesting age by 10 years and a considerable improvement of stem straightness. The third generation will include wood quality traits for different end-uses, including pulp and fibre properties. To facilitate this goal, near infrared spectroscopy (NIRS) was used to estimate chemical composition with regard to lignin, cellulose, hemicelluloses, and extractive content, and lignin quality with regard to 4-hydroxy-phenylpropane/guaiacylpropane ratio. A total of 960 samples were investigated which were collected from a large number of trees (belonging to 80 families obtained by crossing 18 mothers and 20 fathers; there are 12 trees per family). Good calibration data was obtained between NIRS and wet chemistry methods (R2 values higher than 0.9 and good precision of prediction). To complete the NIRS work, kraft cookings in small scale, fibre morphology and microdensitometry investigations were also conducted. Genetic calculations indicated that for a 1% rate of selection on mothers and fathers, genetically induced changes are possible with lignin content (−3.8%), cellulose content (+1.3%), pulp yield (+1.8%), fibre length in pulps (+0.17 mm) and wood density (+50 kg m−3).

Douglas-fir is vulnerable to exceptional and recurrent drought episodes and recovers less well on less fertile sites

ContextSince the 2003 drought and heat wave, there have been many reports of Douglas-fir decline and dieback in France. Given the climate change that is predicted, more frequent drought episodes could induce recurrent decline processes. The nature and background of this threat requires a careful assessment.AimsThe objectives of this study were: (1) to test the hypothesis that the major climate hazard that accounts for the decline in Douglas-fir growth is drought, and (2) to identify the main vulnerability factors involved.MethodsDecline and recovery of radial growth were quantified of Douglas-fir in two particularly affected regions with a dendroecological approach; 899 trees were sampled on 58 plots. Mean climate data, ecological and dendrometric stand characteristics were tested in order to identify potential vulnerability factorsResultsA clear relationship was evidenced between growth and soil water deficit. A severe decline of radial growth was induced by the 2003 drought and was maintained during subsequent years due to recurrent drought episodes. Growth recovery at the stand level was enhanced by soil nutrient fertility.ConclusionRadial growth of Douglas-fir is clearly driven by the intensity of drought. In a context of climate change, an adaptive silvicultural strategy is required in order to cope with more frequent drought events. To mitigate the frequency and the intensity of water shortage episodes, site selection must take into account not only local climate characteristic but also soil properties like maximum extractable water content. Soil nitrogen fertility was found to play a key role for an efficient recovery of radial growth after drought episodes and thus must be preserved.

Wood density proxies of adaptive traits linked with resistance to drought in Douglas fir (Pseudotsuga menziesii (Mirb.) Franco)

AbstractKey messageProxies of adaptive traits for resistance to drought were discovered among original annual ring density variables in Douglas fir.Abstract A comparison of dead and surviving Douglas fir trees following the 2003 drought was made to define proxies of adaptive traits for resistance to drought. Increment cores were sampled from trees from three French regions: Centre, Midi-Pyrénées and Burgundy. Original tree-ring variables were calculated, based on a sliding density criterion dividing the microdensity profile into high- and low-density segments. Tree rings were analysed at each site in a number of consecutive annual rings before the 2003 drought event. Comparison between pairs of surviving and dead trees and between pairs of randomly selected trees (whether dead or alive) supports the evidence of systematic dissimilarities between surviving and dead trees in a number of original density variables. Correlation analysis between original and conventional ring density variables indicates a weak association. We found that the surviving trees were denser than the dead trees in all three sites, but that the denser part of the ring varied from region to region. We identified several original density variables intended to be used as proxies of adaptive traits in future studies of genetic determinism of Douglas fir resistance to drought.

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

Abstract• ContextThe 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.• AimsThis study examined these correlations for different radiata pine genotypes and aimed at detecting potential genotype by climate interactions.• MethodsFour 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.• ResultsCambial 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.• ConclusionClimate 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.