André Rainville

Genomic selection accuracies within and between environments and small breeding groups in white spruce

BackgroundGenomic selection (GS) may improve selection response over conventional pedigree-based selection if markers capture more detailed information than pedigrees in recently domesticated tree species and/or make it more cost effective. Genomic prediction accuracies using 1748 trees and 6932 SNPs representative of as many distinct gene loci were determined for growth and wood traits in white spruce, within and between environments and breeding groups (BG), each with an effective size of Ne ≈ 20. Marker subsets were also tested.ResultsModel fits and/or cross-validation (CV) prediction accuracies for ridge regression (RR) and the least absolute shrinkage and selection operator models approached those of pedigree-based models. With strong relatedness between CV sets, prediction accuracies for RR within environment and BG were high for wood (r = 0.71–0.79) and moderately high for growth (r = 0.52–0.69) traits, in line with trends in heritabilities. For both classes of traits, these accuracies achieved between 83% and 92% of those obtained with phenotypes and pedigree information. Prediction into untested environments remained moderately high for wood (r ≥ 0.61) but dropped significantly for growth (r ≥ 0.24) traits, emphasizing the need to phenotype in all test environments and model genotype-by-environment interactions for growth traits. Removing relatedness between CV sets sharply decreased prediction accuracies for all traits and subpopulations, falling near zero between BGs with no known shared ancestry. For marker subsets, similar patterns were observed but with lower prediction accuracies.ConclusionsGiven the need for high relatedness between CV sets to obtain good prediction accuracies, we recommend to build GS models for prediction within the same breeding population only. Breeding groups could be merged to build genomic prediction models as long as the total effective population size does not exceed 50 individuals in order to obtain high prediction accuracy such as that obtained in the present study. A number of markers limited to a few hundred would not negatively impact prediction accuracies, but these could decrease more rapidly over generations. The most promising short-term approach for genomic selection would likely be the selection of superior individuals within large full-sib families vegetatively propagated to implement multiclonal forestry.

Sources of temporal variation in sap sugar content in a mature sugar maple (Acer saccharum) plantation

Temporal factors affecting sap sugar content (SSC) were assessed in a 65-yr-old plantation of sugar maple (Acer saccharum Marsh.) located in Quebec. A total of 694 maples were measured and the variation in SSC was assessed between 2 sampling years, among 3 days within each sugar season, and among hours within each sampling day. Average individual tree variation in SSC was extensive, ranging from 1.3 to 5.5% for spring 1994, and from 1.4 to 4.8% for spring 1995. The correlation between DBH and SSC was weak for both sampling years. SSC did not vary extensively between sampling hours within days, while it varied significantly among sampling days for both years analysed. However, the largest source of variation in SSC was between the 2 sampling years. Tree relative performances in SSC were moderately conserved among days within years and to a lesser degree, between sampling years. The relative magnitude of the differences between statistical treatments and the stability analysis of tree relative performances indicated that, for adequate estimation of an individual trees SSC, sampling over years appears superior to sampling over several days within a single sugar season. Also, sampling towards the end of the sugar season or during days with lower SSC appears less reliable for mass selection.

The influence of cambial age on breeding for wood properties in Picea glauca

We investigated the influence of cambial age on correlations between different wood traits and the possibility of early selection in order to help decision-making for the improvement of juvenile wood in white spruce (Picea glauca (Moench) Voss). Increment cores were analysed from 375 trees covering 25 open-pollinated families from a 30-year-old provenance–progeny trial in Quebec, Canada. Genetic and phenotypic correlations between different mechanical and fibre anatomy-related wood traits were found to vary with cambial age. Most correlations became stronger in magnitude in rings closer to the bark. An exception is the correlation between microfibril angle (MFA) and the modulus of elasticity where correlations were strongly negative from the pith to the bark. Age–age correlations for different wood traits were found to be high and possible gains from early selection were estimated to be good in ring 8 and older for most traits. MFA was the trait with the strongest potential for selection as early as ring 4, but a detrimental correlation with wood density may represent a drawback of such a juvenile selection approach. Estimates showed that selection concentrated on a few easily measurable traits such as wood density and core length holds promise to obtain superior genetic gains for mechanical properties, but negative impacts would be expected on fibre anatomy traits related to pulp quality. These findings show the need for more carefully planned breeding and selection strategies if one wishes to improve several traits for different end uses.

Genetic Adaptation vs. Ecophysiological Plasticity of Photosynthetic-Related Traits in Young Picea glauca Trees along a Regional Climatic Gradient

Assisted population migration (APM) is the intentional movement of populations within a species range to sites where future environmental conditions are projected to be more conducive to growth. APM has been proposed as a proactive adaptation strategy to maintain forest productivity and to reduce the vulnerability of forest ecosystems to projected climate change. The validity of such a strategy will depend on the adaptation capacity of populations, which can partially be evaluated by the ecophysiological response of different genetic sources along a climatic gradient. This adaptation capacity results from the compromise between (i) the degree of genetic adaptation of seed sources to their environment of origin and (ii) the phenotypic plasticity of functional trait which can make it possible for transferred seed sources to positively respond to new growing conditions. We examined phenotypic variation in morphophysiological traits of six seed sources of white spruce (Picea glauca [Moench] Voss) along a regional climatic gradient in Québec, Canada. Seedlings from the seed sources were planted at three forest sites representing a mean annual temperature (MAT) gradient of 2.2°C. During the second growing season, we measured height growth (H2014) and traits related to resources use efficiency and photosynthetic rate (Amax). All functional traits showed an adaptive response to the climatic gradient. Traits such as H2014, Amax, stomatal conductance (gs), the ratio of mesophyll to stomatal conductance, water use efficiency, and photosynthetic nitrogen-use efficiency showed significant variation in both physiological plasticity due to the planting site and seed source variation related to local genetic adaptation. However, the amplitude of seed source variation was much less than that related to plantation sites in the area investigated. The six seed sources showed a similar level of physiological plasticity. H2014, Amax and gs, but not carboxylation capacity (Vcmax), were correlated and decreased with a reduction of the average temperature of the growing season at seed origin. The clinal variation in H2014 and Amax appeared to be driven by CO2 conductance. The presence of locally adapted functional traits suggests that the use of APM may have advantages for optimizing seed source productivity in future local climates.

Fine-scale geographic variation in photosynthetic-related traits of Picea glauca seedlings indicates local adaptation to climate

Climate-related variations in functional traits of boreal tree species can result both from physiological acclimation and genetic adaptation of local populations to their biophysical environment. To improve our understanding and prediction of the physiological and growth responses of populations to climate change, we studied the role of climate of seed origin in determining variations in functional traits and its implications for tree improvement programs for a commonly reforested boreal conifer, white spruce (Picea glauca (Moench) Voss). We evaluated growth, root-to-shoot ratio (R/S), specific leaf area (SLA), needle nitrogen (N(mass)), total non-structural carbohydrates (NSC) and photosynthetic traits of 3-year-old seedlings in a greenhouse experiment using seed from six seed orchards (SO) representing the different regions where white spruce is reforested in Québec. Height and total dry mass (TDM) were positively correlated with photosynthetic capacity (A(max)), stomatal conductance (g(s)) and mesophyll conductance (g(m)). Total dry mass, but not height growth, was strongly correlated with latitude of seed origin (SO) and associated climate variables. A(max), g(s), g(m) and more marginally, photosynthetic nitrogen-use efficiency (PNUE) were positively associated with the mean July temperature of the SO, while water use efficiency (WUE) was negatively associated. Maximum rates of carboxylation (V(cmax)), maximum rates of electron transport (J(max)), SLA, N(mass), NSC and R/S showed no pattern. Our results did not demonstrate a higher Amax for northern seed orchards, although this has been previously hypothesized as an adaptation mechanism for maintaining carbon uptake in northern regions. We suggest that gs, gm, WUE and PNUE are the functional traits most associated with fine-scale geographic clines and with the degree of local adaptation of white spruce populations to their biophysical environments. These geographic patterns may reflect in situ adaptive genetic differences in photosynthetic efficiency along the cline.

Genetic Control and Nursery-Plantation Genotypic Correlations for Growth Characteristics of White Spruce Somatic Clones

Height and diameter of white spruce clones were measured over two growing seasons in a forest nursery and for 4 more yr at two field sites. Clonal heritability (Hc 2) was moderate and stable in the nursery, but decreased with age, reaching 0.26 and 0.11 for height and 0.14 and 0.04 for diameter, respectively, at the two field sites. Height exhibited weak to moderate Hc 2 and strong genotypic correlations with the different growth characteristics. The strong age-age genotypic correlations (≥0.809) and nursery-plantation correlations (≥0.347) for height indicate that selection for height can be conducted at a young age thus reducing the costs of clonal tests.

Morpho-physiological variation of white spruce seedlings from various seed sources and implications for deployment under climate change

Because of changes in climatic conditions, tree seeds originating from breeding programs may no longer be suited to sites where they are currently sent. As a consequence, new seed zones may have to be delineated. Assisted migration consists of transferring seed sources that match the future climatic conditions to which they are currently adapted. It represents a strategy that could be used to mitigate the potential negative consequences of climate change on forest productivity. Decisions with regard to the choice of the most appropriate seed sources have to rely on appropriate knowledge of morpho-physiological responses of trees. To meet this goal, white spruce (Picea glauca [Moench] Voss) seedlings from eight seed orchards were evaluated during two years in a forest nursery, and at the end of the first growing season on three plantation sites located in different bioclimatic domains in Quebec. The morpho-physiological responses obtained at the end of the second growing season (2+0) in the nursery made it possible to cluster the orchards into three distinct groups. Modeling growth curves of these different groups showed that the height growth of seedlings from the second-generation and southern first-generation seed orchards was significantly higher than that of those from other orchards, by at least 6%. A multiple regression model with three climatic variables (average growing season temperature, average July temperature, length of the growing season) showed that the final height of seedlings (2+0) from the first-generation seed orchards was significantly related to the local climatic conditions at the orchard sites of origin where parental trees from surrounding natural populations were sampled to provide grafts for orchard establishment. Seedling height growth was significantly affected by both seed source origins and planting sites, but the relative ranking of the different seed sources was maintained regardless of reforestation site. This knowledge could be used, in conjunction with transfer models, to refine operational seed transfer rules and select the most suitable sites in an assisted migration strategy.

Genetic control and evolutionary potential of a constitutive resistance mechanism against the spruce budworm ( Choristoneura fumiferana ) in white spruce ( Picea glauca )

Insect herbivory may drive evolution by selecting for trees with heritable resistance against defoliation. The spruce budworm (Choristoneura fumiferana, SBW) is a highly damaging forest insect pest that can affect population structure of white spruce (Picea glauca) in North America. Resistance against SBW was recently described in white spruce and was linked to three constitutive resistance biomarkers: the phenolic compounds piceol and pungenol, and expression of a beta-glucosidase encoding gene (Pgβglu-1). We investigated the phenotypic variability and heritability of these resistance biomarkers and of picein, the precursor of piceol, in the foliage of 874 trees belonging to 33 full-sib families and 71 clonal lines under evaluation in seven field locations in Eastern Canada. We aimed to (i) determine their genetic control, (ii) estimate the genetic and phenotypic correlations among defense biomarkers, and (iii) determine whether their constitutive levels are associated with detrimental trade-offs on growth. Quantitative genetics analyses indicated that all four traits are moderately to highly heritable. The full-sib and clonal analyses showed that additive and non-additive genetic effects play major and minor roles, respectively. Positive genetic and phenotypic correlations between resistance biomarkers and primary growth indicated that there is no trade-off between total height and height increment and resistance traits, contradicting the GDBH (Growth Differentiation Balance Hypothesis). Our findings about the predominant additive genetic basis of the resistance biomarkers show that adaptive evolution of white spruce natural populations to resist to SBW is possible and that potentially important gains could also be expected from artificial selection.

Thermal acclimation of photosynthesis and respiration of southern and northern white spruce seed sources tested along a regional climatic gradient indicates limited potential to cope with temperature warming

Abstract Background and Aims Knowledge of thermal acclimation of physiological processes of boreal tree species is necessary to determine their ability to adapt to predicted global warming and reduce the uncertainty around the anticipated feedbacks of forest ecosystems and global carbon cycle to climate change. The objective of this work was to examine the extent of thermal acclimation of net photosynthesis (An) and dark respiration (Rd) of two distant white spruce (Picea glauca) seed sources (from south and north of the commerial forest zone in Québec) in response to latitudinal and seasonal variations in growing conditions. Methods The temperature responses of An, its biochemical and biophysical limitations, and Rd were measured in 1-year-old needles of seedlings from the seed sources growing in eight forest plantations along a regional thermal gradient of 5.5 °C in Québec, Canada. Key Results The average optimum temperature (Topt) for An was 19 ± 1.2 °C and was similar among seed sources and plantation sites along the thermal gradient. Net photosynthesis at Topt (Aopt) varied significantly among plantation sites and was quadratically related to the mean July temperature (MJT) of plantation sites. Topt for mesophyll conductance, maximum electron transport rate and maximum rate of carboxylation were 28, 22 and 30 °C, respectively. Basal respiration rate (Rd at 10 °C) was linearly and negatively associated with MJT. Q10 of Rd (the rate of change in Rd with a 10 °C increase in temperature) did not show any significant relationship with MJT and averaged 1.5 ± 0.1. The two seed sources were similar in their thermal responses to latitudinal and seasonal variations in growing conditions. Conclusions The results showed moderate thermal acclimation of respiration and no evidence for thermal acclimation of photosynthesis or local genetic adaptation for traits related to thermal acclimation. Therefore, growth of local white spruces may decline in future climates.

Family Variation in the Morphology and Physiology of White Spruce (Picea glauca) Seedlings in Response to Elevated CO2 and Temperature

Tree improvement programs aim to develop families that are well-adapted to future growing conditions. To gain insight into the stability of the family genetic response to climate change, white spruce (Picea glauca) seedlings from 60 full-sib families were subjected to a combination of two temperature regimes and two levels of CO2 over two growing seasons. There was positive effect of warmer temperatures and higher CO2 on some growth variables but no significant family × treatment interactions. Instantaneous water use efficiency was the only physiological trait that was affected positively by the CO2 treatment, showing a 51% increase that was consistent across families.