Mapping the genetic architecture of forest tree traits is important in order to understand the evolutionary forces that have shaped these traits and to facilitate the development of genomic-based breeding strategies. We examined the number, size, and distribution of allelic effects influencing eight types of traits using 30 published mapping studies (linkage and association mapping) in forest trees. The sizes of allelic effects, measured as the phenotypic variance explained, generally showed a severely right-skewed distribution. We estimated the numbers of underlying causal effects (n _qtl ) for different trait categories by improving a method previously developed by Otto and Jones (Genetics 156:2093–2107, 2000). Estimates of n _qtl based on association mapping studies were generally higher (median at 643) than those based on linkage mapping (median at 33). Comparisons with simulated linkage and association mapping data suggested that the lower n _qtl estimates for the linkage mapping studies could partly be explained by fewer causal loci segregating within the full-sib family populations normally used, but also by the cosegregation of causal loci due to limited recombination. Disease resistance estimates based on linkage mapping studies had the lowest median of four underlying effects, while growth traits based on association mapping had about 580 effects. Theoretically, the capture of 50% of the genetic variation would thus require a population size of about 200 for disease resistance in linkage mapping, while growth traits in association mapping would require about 25,000. The adequacy and reliability of the improved method was successfully verified by applying it to the simulated data.

Key message: There is considerable genetic gain of tree volume from clonal deployment in Norway spruce (Picea abies(L.) H. Karst) and clonal deployment will have at least 50% more or double genetic gain than the seedling deployment. Context: Genetic parameters and genetic gains for wood quality and growth traits were estimated in six large clonal progeny trials. Aims: Develop the optimal clonal deployment strategy of Norway spruce in Sweden. Methods: Wood quality and growth traits were measured in all clonal trials and additive and non-additive genetic variances are partitioned. Results: Additive and non-additive genetic variances were equally important for growth traits while non-additive variance was small or not significant for wood quality trait. The genetic gain predicted for clonal deployment was greater than any of the other four deployment strategies. Selecting the top 1% of tested clones (clonal forestry) would have 48.4% and 134.6% more gain than the gain predicted for the seedling deployment of selected full-sib families and half-sib family (family forestry), respectively, at the same selection intensity. Conclusion: This study highlights that testing of 30–40 clones per family would maximize the realized genetic gain for different clonal selection scenarios, either selecting the best ten or 20 clones without any co-ancestry restrictions or selecting the best single clone from each of the best ten or 20 families (e.g., co-ancestry restriction). Clonal mean selection and vegetative deployment are the most effective.

Willow species (Salix) are important as short‐rotation biomass crops for bioenergy, which creates a demand for faster genetic improvement and breeding through deployment of molecular marker‐assisted selection (MAS). To find markers associated with important adaptive traits, such as growth and phenology, for use in MAS, we genetically dissected the trait variation of a Salix viminalis (L.) population of 323 accessions. The accessions were sampled throughout northern Europe and were established at two field sites in Pustnäs, Sweden, and at Woburn, UK, offering the opportunity to assess the impact of genotype‐by‐environment interactions (G × E) on trait–marker associations. Field measurements were recorded for growth and phenology traits. The accessions were genotyped using 1536 SNP markers developed from phenology candidate genes and from genes previously observed to be differentially expressed in contrasting environments. Association mapping between 1233 of these SNPs and the measured traits was performed taking into account population structure and threshold selection bias. At a false discovery rate (FDR) of 0.2, 29 SNPs were associated with bud burst, leaf senescence, number of shoots or shoot diameter. The percentage of accession variation () explained by these associations ranged from 0.3% to 4.4%, suggesting that the studied traits are controlled by many loci of limited individual impact. Despite this, a SNP in the EARLY FLOWERING 3 gene was repeatedly associated (FDR < 0.2) with bud burst. The rare homozygous genotype exhibited 0.4–1.0 lower bud burst scores than the other genotype classes on a five‐grade scale. Consequently, this marker could be promising for use in MAS and the gene deserves further study. Otherwise, associations were less consistent across sites, likely due to their small estimates and to considerable G × E interactions indicated by multivariate association analyses and modest trait accession correlations across sites (0.32–0.61).

Biased sex ratios are common among dioecious plant species despite the theoretical prediction of selective advantage of even sex ratios. Albeit the high prevalence of deviations from even sex ratios, the genetic causes to sex biases are rarely known outside of a few model species. Here we present a mechanism underlying the female biased sex ratio in the dioecious willow species Salix viminalis. We compared the segregation pattern of genome-wide single nucleotide polymorphism markers in two contrasting bi-parental pedigree populations, the S3 with even sex ratio and the S5 with a female biased sex ratio. With the segregation analysis and comparison between the two populations, we were able to demonstrate that sex determination and sex ratio distortion are controlled by different genetic mechanisms. We furthermore located the sex ratio distorter locus to a Z/W-gametologous region on chromosome 15, which was in close linkage with the sex determination locus. Interestingly, all males in the population with biased sex ratio have in this sex ratio distorter locus the same genotype, meaning that males with the Z1/Z3-genotype were missing from the population, thereby creating the 2:1 female biased sex ratio. We attribute the absence of Z1/Z3 males to an allelic incompatibility between maternally and paternally inherited alleles in this sex ratio distorter locus. Due to the tight linkage with the sex determination locus only male individuals are purged from the population at an early age, presumably before or during seed development. We showed that such allelic incompatibility could be stably maintained over evolutionary times through a system of overdominant or pseudooverdominant alleles. Thus, it is possible that the same mechanism generates the female biased sex ratio in natural willow populations.

Salix spp. are high-productivity crops potentially used for lignocellulosic biofuels such as bioethanol. In general, pretreatment is needed to facilitate the enzymatic depolymerization process. Biomass resistance to degradation, i.e., biomass recalcitrance, is a trait which can be assessed by measuring the sugar released after combined pretreatment and enzymatic hydrolysis. We have examined genetic parameters of enzymatic sugar release and other traits related to biorefinery use in a population of 286 natural Salix viminalis clones. Furthermore, we have evaluated phenotypic and genetic correlations between these traits and performed a genomewide association mapping analysis using a set of 19,411 markers. Sugar release (glucose and xylose) after pretreatment and enzymatic saccharification proved highly variable with large genetic and phenotypic variations, and chip heritability estimates (h2) of 0.23–0.29. Lignin syringyl/guaiacyl (S/G) ratio and wood density were the most heritable traits (h2 = 0.42 and 0.59, respectively). Sugar release traits were positively correlated, phenotypically and genetically, with biomass yield and lignin S/G ratio. Association mapping revealed seven marker–trait associations below a suggestive significance threshold, including one marker associated with glucose release. We identified lignin S/G ratio and shoot diameter as heritable traits that could be relatively easily evaluated by breeders, making them suitable proxy traits for developing low-recalcitrance varieties. One marker below the suggestive threshold for marker associations was identified for sugar release, meriting further investigation while also highlighting the difficulties in employing genomewide association mapping for complex traits.

In breeding Norway spruce, selection for improved growth and survival is performed at age 10–15 years in order to optimize genetic gain per year. We investigated whether a selection based on wood traits such as density and grain angle, measured under bark in the field at the same age would be informative enough with respect to structural quality traits of sawn boards. To achieve this objective, a sawing study was conducted on the butt logs of 401 trees from a 34-year-old Norway spruce progeny trial situated in southern Sweden. Stem discs were excised from the top of the logs and radial profile data of grain angle, and wood density was recorded for specific annual rings. The sawn and dried boards were assessed for structural traits such as twist, board density, bending stiffness (static modulus of elasticity, sMoE) and bending strength (modulus of rupture, MoR). Additive genetic correlations (r_a) between single annual ring density measurements and board density, sMoE and MoR were consistently strong (r_a> 0.7) for annual rings 5–13. Genetic correlations of similar magnitude between grain angle and board twist were estimated for all investigated annual rings (from 2 to around 26 under bark). Consequently, it was found that indirect selection for wood density and grain angle at the tree age 10–16 years would result in more genetic gain per year than selection at later ages. This makes it feasible to perform simultaneous selection of progeny in the field for both growth and wood traits at similar ages.