Summary
Norway spruce is a boreal forest tree species of significant ecological and economic importance. Hence there is a strong imperative to dissect the genetics underlying important wood quality ...traits in the species. We performed a functional genome‐wide association study (GWAS) of 17 wood traits in Norway spruce using 178 101 single nucleotide polymorphisms (SNPs) generated from exome genotyping of 517 mother trees. The wood traits were defined using functional modelling of wood properties across annual growth rings. We applied a Least Absolute Shrinkage and Selection Operator (LASSO‐based) association mapping method using a functional multilocus mapping approach that utilizes latent traits, with a stability selection probability method as the hypothesis testing approach to determine a significant quantitative trait locus. The analysis provided 52 significant SNPs from 39 candidate genes, including genes previously implicated in wood formation and tree growth in spruce and other species. Our study represents a multilocus GWAS for complex wood traits in Norway spruce. The results advance our understanding of the genetics influencing wood traits and identifies candidate genes for future functional studies.
Significance Statement
Wood provides both structural support and a transport route for water and solutes in trees. Our work provides a framework to dissect the genetic nature of wood formation and adds to our understanding of tree growth and development. With the current research focus on wood cell wall biosynthesis in general, and lignocellulose feedstock for biorefineries, we believe that this contribution will be of wide interest for the plant science community.
Xylan is one of the main compounds determining wood properties in hardwood species. The xylan backbone is thought to be synthesized by a synthase complex comprising two members of the GT43 family. We ...downregulated all GT43 genes in hybrid aspen (Populus tremula × tremuloides) to understand their involvement in xylan biosynthesis.
All three clades of the GT43 family were targeted for downregulation using RNA interference individually or in different combinations, either constitutively or specifically in developing wood.
Simultaneous downregulation in developing wood of the B (IRX9) and C (IRX14) clades resulted in reduced xylan Xyl content relative to reducing end sequence, supporting their role in xylan backbone biosynthesis. This was accompanied by a higher lignocellulose saccharification efficiency. Unexpectedly, GT43 suppression in developing wood led to an overall growth stimulation, xylem cell wall thinning and a shift in cellulose orientation. Transcriptome profiling of these transgenic lines indicated that cell cycling was stimulated and secondary wall biosynthesis was repressed. We suggest that the reduced xylan elongation is sensed by the cell wall integrity surveying mechanism in developing wood.
Our results show that wood-specific suppression of xylan-biosynthetic GT43 genes activates signaling responses, leading to increased growth and improved lignocellulose saccharification.
Annual growth, fibre and wood properties of Norway spruce are all under strong influence from genetics, age and weather. They change dynamically, particularly at young ages. Most genetic research and ...tree improvement programs are based on data from this most dynamic phase of the life of trees, affected by differences in weather among sites and years. In the work presented, influences of age and weather were investigated and modelled at the detail of annual rings and at the sub-tree ring level of earlywood, transitionwood and latewood. The data used were analysed from increment cores sampled at age 21 years from almost 6000 Norway spruce trees of known genetic origin, grown on two sites in southern Sweden. The traits under investigation were radial growth, cell widths, cell numbers, cell wall thickness and coarseness as a measure of biomass allocation at cell level. General additive mixed models (GAMMs) were fitted to model the influences of age, local temperature and precipitation. The best models were obtained for number of tracheids formed per year, ring width, average radial tracheid width in earlywood, and ring averages for tangential tracheid width and coarseness. Considering the many sources behind the huge variation, the explained part of the variability was high. For all traits, models were developed using both total tree age and cambial age (ring number) to express age. Comparisons indicate that the number of cell divisions and ring width are under stronger control of tree age, but the other traits under stronger control of cambial age. The models provide a basis to refine data prior to genetic evaluations by compensating for estimated differences between sites and years related to age and weather rather than genetics. Other expected applications are to predict performance of genotypes in relation to site or climate and simulation of climate change scenarios.
We studied the interactive effects of elevated concentrations of CO2 and O3 on radial growth and wood properties of four trembling aspen (Populus tremuloides Michx.) clones and paper birch (Betula ...papyrifera Marsh.) saplings. The material for the study was collected from the Aspen FACE (free‐air CO2 enrichment) experiment in Rhinelander (WI, USA). Trees had been exposed to four treatments control, elevated CO2 (560 ppm), elevated O3 (1.5 times ambient) and combined CO2 + O3 during growing seasons 1998–2008. Most treatment responses were observed in the early phase of experiment. Our results show that the CO2‐ and O3‐exposed aspen trees displayed a differential balance between efficiency and safety of water transport. Under elevated CO2, radial growth was enhanced and the trees had fewer but hydraulically more efficient larger diameter vessels. In contrast, elevated O3 decreased radial growth and the diameters of vessels and fibres. Clone‐specific decrease in wood density and cell wall thickness was observed under elevated CO2. In birch, the treatments had no major impacts on wood anatomy or wood density. Our study indicates that short‐term impact studies conducted with young seedlings may not give a realistic view of long‐term ecosystem responses.
We studied whether growth changes induced by elevated CO2 and/or O3 were maintained during 11‐year exposure in the field and how they reflected to wood structure of aspen and birch. Neither CO2 nor O3 responses were consistent throughout the experiment, but they both varied annually and were more often seen early in the experiment. Our results show that the CO2‐ and O3‐exposed aspen trees displayed a differential balance between efficiency and safety of water transport: the CO2 trees with enhanced radial growth had fewer but hydraulically more efficient larger‐diameter vessels. Our study indicates that short‐term impact studies conducted with young seedlings may not give a realistic view of long‐term ecosystem responses.
Genomic selection (GS) or genomic prediction is considered as a promising approach to accelerate tree breeding and increase genetic gain by shortening breeding cycle, but the efforts to develop ...routines for operational breeding are so far limited. We investigated the predictive ability (PA) of GS based on 484 progeny trees from 62 half-sib families in Norway spruce (Picea abies (L.) Karst.) for wood density, modulus of elasticity (MOE) and microfibril angle (MFA) measured with SilviScan, as well as for measurements on standing trees by Pilodyn and Hitman instruments.
GS predictive abilities were comparable with those based on pedigree-based prediction. Marker-based PAs were generally 25-30% higher for traits density, MFA and MOE measured with SilviScan than for their respective standing tree-based method which measured with Pilodyn and Hitman. Prediction accuracy (PC) of the standing tree-based methods were similar or even higher than increment core-based method. 78-95% of the maximal PAs of density, MFA and MOE obtained from coring to the pith at high age were reached by using data possible to obtain by drilling 3-5 rings towards the pith at tree age 10-12.
This study indicates standing tree-based measurements is a cost-effective alternative method for GS. PA of GS methods were comparable with those pedigree-based prediction. The highest PAs were reached with at least 80-90% of the dataset used as training set. Selection for trait density could be conducted at an earlier age than for MFA and MOE. Operational breeding can also be optimized by training the model at an earlier age or using 3 to 5 outermost rings at tree age 10 to 12 years, thereby shortening the cycle and reducing the impact on the tree.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The current distribution and population structure of many species were, to a large extent, shaped by cycles of isolation in glacial refugia and subsequent population expansions. Isolation in and ...postglacial expansion through heterogeneous environments led to either neutral or adaptive divergence. Norway spruce is no exception, and its current distribution is the consequence of a constant interplay between evolutionary and demographic processes. We investigated population differentiation and adaptation of Norway spruce for juvenile growth, diameter of the stem, wood density, and tracheid traits at breast height. Data from 4461 phenotyped and genotyped Norway spruce from 396 half‐sib families in two progeny tests were used to test for divergent selection in the framework of QST vs. FST. We show that the macroscopic resultant trait (stem diameter), unlike its microscopic components (tracheid dimensions) and juvenile growth, was under divergent selection that predated the Last Glacial Maximum. Altogether, the current variation in these phenotypic traits in Norway spruce is better explained by local adaptation to ancestral environments than to current ones, where populations were partly preadapted, mainly through growth‐related traits.
Through the use of genome-wide association studies (GWAS) mapping it is possible to establish the genetic basis of phenotypic trait variation. Our GWAS study presents the first such effort in Norway ...spruce (Picea abies (L). Karst.) for the traits related to wood tracheid characteristics. The study employed an exome capture genotyping approach that generated 178 101 Single Nucleotide Polymorphisms (SNPs) from 40 018 probes within a population of 517 Norway spruce mother trees. We applied a least absolute shrinkage and selection operator (LASSO) based association mapping method using a functional multi-locus mapping approach, with a stability selection probability method as the hypothesis testing approach to determine significant Quantitative Trait Loci (QTLs). The analysis has provided 30 significant associations, the majority of which show specific expression in wood-forming tissues or high ubiquitous expression, potentially controlling tracheids dimensions, their cell wall thickness and microfibril angle. Among the most promising candidates based on our results and prior information for other species are: Picea abies BIG GRAIN 2 (PabBG2) with a predicted function in auxin transport and sensitivity, and MA_373300g0010 encoding a protein similar to wall-associated receptor kinases, which were both associated with cell wall thickness. The results demonstrate feasibility of GWAS to identify novel candidate genes controlling industrially-relevant tracheid traits in Norway spruce.
Unfavorable genetic correlations between growth and wood quality traits are one of the biggest challenges in advanced conifer breeding programs. To examine and deal with such correlation, increment ...cores were sampled at breast height from 5,618 trees in 524 open-pollinated families in two 21-year-old Norway spruce progeny trials in southern Sweden, and age trends of genetic variation, genetic correlation, and efficiency of selection were investigated. Wood quality traits were measured on 12-mm increment cores using SilviScan. Heritability was moderate (~0.4–0.5) for wood density and modulus of elasticity (MOE) but low (~0.2) for microfibril angle (MFA). Different age trends were observed for wood density, MFA, and MOE, and the lower heritability of MFA relative to wood density and MOE in Norway spruce contrasted with general trends of the three wood quality traits in pine. Genetic correlations among growth, wood density, MFA, and MOE increased to a considerably high value from pith to bark with unfavorable genetic correlations (−0.6 between growth and wood density, −0.74 between growth and MOE). Age–age genetic correlations reached 0.9 after ring 4 for diameter at breast height (DBH), wood density, MFA, and MOE traits. Early selections at ring 10 for diameter and at ring 6 or 7 for wood quality traits had similar effectiveness as selection conducted at reference ring 15. Selection based on diameter alone produced 19.0 % genetic gain in diameter but resulted in 4.8 % decrease in wood density, 9.4 % decrease in MOE, and 8.0 % increase in MFA. Index selection with a restriction of no change in wood density, MOE, and MFA, respectively, produced relatively lower genetic gains in diameter (16.4, 12.2, and 14.1 %, respectively), indicating such index selection could be implemented to maintain current wood density. Index selection using economic weights is, however, recommended for maximum economic efficiency.
The objective of the study was to investigate the interactive effects of elevated atmospheric carbon dioxide concentration, CO₂, and temperature on the wood properties of mature field-grown Norway ...spruce (Picea abies (L.) Karst.) trees. Material for the study was obtained from an experiment in Flakaliden, northern Sweden, where trees were grown for 3 years in whole-tree chambers at ambient (365 μmol mol⁻¹) or elevated CO₂ (700 μmol mol⁻¹) and ambient or elevated air temperature (ambient +5.6 °C in winter and ambient +2.8 °C in summer). Elevated temperature affected both wood chemical composition and structure, but had no effect on stem radial growth. Elevated temperature decreased the concentrations of acetone-soluble extractives and soluble sugars, while mean and earlywood (EW) cell wall thickness and wood density were increased. Elevated CO₂ had no effect on stem wood chemistry or radial growth. In wood structure, elevated CO₂ decreased EW cell wall thickness and increased tracheid radial diameter in latewood (LW). Some significant interactions between elevated CO₂ and temperature were found in the anatomical and physical properties of stem wood (e.g. microfibril angle, and LW cell wall thickness and density). Our results show that the wood material properties of mature Norway spruce were altered under exposure to elevated CO₂ and temperature, although stem radial growth was not affected by the treatments.
Tracheid length and width patterns from pith to bark at a height of 0.6 m in uneven-aged Norway spruce (Picea abies (L.) Karst.) trees were studied. Identification of the main factors and a ...comparison with even-aged stands were also carried out. Ninety-six trees were sampled from experimental stands in southern Finland. The material encompassed the variation in tracheid properties from early years to silvicultural maturity, i.e., from corewood to outerwood up to a cambial age of 111 years. Data from 39 Norway spruce trees from even-aged stands were utilized for comparison. Models fitted to the data indicated that annual ring widths did not influence mean tracheid dimensions but that the latewood proportion had a significant influence on tracheid dimensions. Tracheids in uneven-aged stands were slightly wider and longer at the base of the stem in trees with a similar diameter, cambial age, and annual ring number.
Celotno besedilo
Dostopno za:
BF, DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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