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Aim: Seedling recruitment is a vital process for forest regeneration and is influenced by various factors such as stand composition, climate, and soil disturbance. We conducted a long‐term field experiment (18 years) to study the effects of these factors and their interactions on seedling recruitment. Location: Our study focused on five main species in boreal mixed woods of eastern Canada: trembling aspen (Populus tremuloides), paper birch (Betula papyrifera), white spruce (Picea glauca), balsam fir (Abies balsamea), and white cedar (Thuja occidentalis). Methods: Sixteen 1‐m2 seedling monitoring subplots were set up in each of seven stands originating from different wildfires (fire years ranging from 1760 to 1944), with a soil scarification treatment applied to every other subplot. Annual new seedling counts were related to growing‐season climate (mean temperature, growing degree days and drought code), scarification, and stand effects via a Bayesian generalized linear mixed model. Results: Soil scarification had a large positive effect on seedling recruitment for three species (aspen, birch and spruce). As expected, high mean temperatures during the seed production period (two years prior to seedling emergence) increased seedling recruitment for all species but aspen. Contrary to other studies, we did not find a positive effect of dry conditions during the seed production period. Furthermore, high values of growing degree days suppressed conifer seedling recruitment. Except for white cedar, basal area was weakly correlated with seedling abundance, suggesting a small number of reproductive individuals is sufficient to saturate seedling recruitment. Conclusion: Our findings underscore the importance of considering multiple factors, such as soil disturbance, climate, and stand composition, as well as their effects on different life stages when developing effective forest management strategies to promote regeneration in boreal mixed‐wood ecosystems. [ABSTRACT FROM AUTHOR]

Data collected over a 30-year period from an experiment replicated across 21 locations in western Canada are used to explore the effects of precommercial thinning of trembling aspen to a range of densities in combination with three initial white spruce densities on tree growth and stand dynamics. Increasing differentiation amongst the 15 treatments was observed with age after thinning for both spruce and aspen responses. Spruce height and diameter declined with increasing aspen density. At age 10 spruce diameter with no aspen was 1.5× that of spruce in unthinned while it was 2.6× that of spruce in unthinned at age 28. Following thinning aspen DBH and crown width declined with increasing density of retained aspen while slenderness and height to crown base of aspen increased. Thinning to 1500 aspen·ha−1 resulted in aspen DBH that was 22% larger relative to the unthinned at age 28. Spruce volume and stemwood biomass at age 90, estimated using the Mixedwood Growth Model (MGM21), declined with increasing initial aspen density, with the complete removal of aspen resulting in nearly double the spruce volume of unthinned plots. However, total stand volume, total stand biomass, and stemwood carbon at age 90 are predicted to be largest in mixed stands with aspen densities of 1500 stems·ha−1 or higher. [ABSTRACT FROM AUTHOR]

Rapid establishment of closed canopy cover is a key goal in land reclamation. While re-establishment of understory vegetation is likely to positively contribute to this goal, native herbaceous species are not typically planted on reclaimed sites though it is an alternative to reliance on natural regeneration. The objective of the present study was to test the principle of hitchhiker production of a tree species with native herbaceous species on recently reclaimed industrial sites with the aim of (1) understanding growth tradeoffs when mixing these species in the same container and (2) if hitchhiking had negative growth and survival outcomes for white spruce (Picea glauca) seedlings compared with singly grown white spruce seedlings. White spruce seeds were sown in nursery containers of contrasting cavity sizes, followed by sowing a forb species (Eurybia conspicua or Chamerion angustifolium) at different time intervals or grown singly. Larger cavities and earlier sow dates were associated with higher persistence and growth of the forbs, while later sow dates were associated with larger white spruce seedlings. We observed no evidence to suggest that hitchhiked seedlings were at a competitive disadvantage in terms of growth or survival compared with singly grown seedlings that were initially similar in size. [ABSTRACT FROM AUTHOR]

The utilization of genotyping has gained significant popularity in tree improvement programs, aiding in enhancing the precision of breeding values, removing pedigree errors, the assessment of genetic diversity, and evaluating pollen contamination. Our study explores the impact of utilizing 5308 SNP markers to genotype seed orchard parents (166), progeny in progeny trials (667), and seedlot orchard seedlings (780), to simultaneously enhance variance components, breeding values, genetic diversity estimates, and pollen flow in the Region I white spruce (Picea glauca) breeding program in central Alberta (Canada). We compared different individual tree mixed models, including pedigree-based (ABLUP), genomic-based (GBLUP), and single-step pedigree-genomic-based (ssGBLUP) models, to estimate variance components and predict breeding values for the height and diameter at breast height traits. The highest heritability estimates were achieved using the ssGBLUP approach, resulting in improved breeding value accuracy compared to the ABLUP and GBLUP models for the studied growth traits. In the six orchard seedlots tested, the genetic diversity of the seedlings remained stable, characterized by an average of approximately 2.00 alleles per SNP, a Shannon Index of approximately 0.44, and an expected and observed heterozygosity of approximately 0.29. The pedigree reconstruction of seed orchard seedlings successfully identified consistent parental contributions and equal genotype contributions in different years. Pollen contamination levels varied between 11% and 70% using SNP markers and 8% to 81% using pollen traps, with traps both over- and under-estimating contamination. Overall, integrating genomic information from parents and offspring empowers forest geneticists and breeders in the Region I white spruce breeding program to correct errors, conduct backward and forward selections with greater precision, gain a deeper understanding of the orchard's genetic structure, select superior seedlots, and accurately estimate the genetic worth of each orchard lot, which can ultimately result in increased and more precise estimates of genetic gain in the studied growth traits. [ABSTRACT FROM AUTHOR]

Data collected over a 30-year period from an experiment replicated across 21 locations in western Canada are used to explore the effects of precommercial thinning of trembling aspen to a range of densities in combination with three initial white spruce densities on tree growth and stand dynamics. Increasing differentiation amongst the 15 treatments was observed with age after thinning for both spruce and aspen responses. Spruce height and diameter declined with increasing aspen density. At age 10 spruce diameter with no aspen was 1.5× that of spruce in unthinned while it was 2.6× that of spruce in unthinned at age 28. Following thinning aspen DBH and crown width declined with increasing density of retained aspen while slenderness and height to crown base of aspen increased. Thinning to 1500 aspen·ha−1 resulted in aspen DBH that was 22% larger relative to the unthinned at age 28. Spruce volume and stemwood biomass at age 90, estimated using the Mixedwood Growth Model (MGM21), declined with increasing initial aspen density, with the complete removal of aspen resulting in nearly double the spruce volume of unthinned plots. However, total stand volume, total stand biomass, and stemwood carbon at age 90 are predicted to be largest in mixed stands with aspen densities of 1500 stems·ha−1 or higher.

Much is still unknown about the growth and physiological responses of trees to global change at the northern treeline. We combined tree‐ring width data with century‐long stable carbon and oxygen isotope records to investigate growth and physiological responses of white spruce at two treeline sites in the Canadian Arctic to concurrent increases in temperature, atmospheric CO2 concentration (ca), and decline in sea ice extent over the past century. The tree‐ring records were assessed during three periods with contrasting climatic conditions: (a) the early 20th century warming, (b) the 1940–1970 cooling period, and (c) the anthropogenic late 20th century warming period. We found opposing growth trends between the two sites, but similar carbon isotope discrimination (Δ13C) and intrinsic water‐use efficiency (iWUE) trajectories. While tree growth (defined as basal area increment) increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, growth declined after 1950 at the more interior site. At both sites, Δ13C slightly increased over these periods. However, trees showed a nonlinear response to increased ca, shifting after 1970 from a passive stomatal response (i.e., no changes in iWUE) to an active response (i.e., a moderate ∼12% increase in iWUE). Further, our isotope‐based findings do not support the idea that temperature‐induced drought stress caused the divergent growth trends at our treeline sites. This study thus highlights nonlinear and complex physiological and growth adjustments to concomitant changes in temperature, sea ice extent, and ca over the last century at the northern treeline. Plain Language Summary: In the last decades, Arctic ecosystems have experienced unprecedented warmer temperatures, higher atmospheric carbon dioxide concentrations, and a decline in sea ice extent. However, it is still unknown how these changes in environmental factors have impacted tree populations at their northern distribution limits. Here, we examine how the growth and water‐use efficiency of white spruce trees have changed over the last century in response to warmer temperatures, higher carbon dioxide concentrations, and sea ice decline at two treeline sites in the Canadian Arctic. We found that tree growth increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, while growth declined after 1950 at the more interior site. At both sites, the water‐use efficiency of trees was stable until 1970. Following the rapid temperature and carbon dioxide increase since the 1970s, trees have become more efficient in using water. The contrasting growth trends found at our study sites highlight complex growth and physiological adjustments of white spruce trees to rapid and simultaneous changes in temperature, sea ice extent, and carbon dioxide concentrations in the Arctic. Key Points: Complex growth and physiological responses of white spruce trees to changes in temperature and sea ice extent inferred from stable isotopesWhite spruce radial growth at Arctic latitudinal treeline decreased at an interior site and increased at a site closer to the Arctic OceanWhite spruce shifted from a passive to an active water‐use efficiency response to rising CO2 concentration over the 20th century [ABSTRACT FROM AUTHOR]

Plants require metals such as copper to survive, but an excess of metals can have phytotoxic effects. The objectives of this study were to (1) analyze the expression of genes associated with copper resistance (RAN1 and MRP4) and (2) assess the effects of copper and potassium ions on global DNA methylation in white spruce (P. glauca). Seedlings were treated with three concentrations of copper sulfate including 1312 mg/kg, 656 mg/kg, and 130 mg/kg. Potassium sulfate and water were used as controls. DNA and RNA were extracted from roots and needles. The levels of gene expression were measured using RT-qPCR and the global 5-methyl cytosine was assessed using the Abcam ELISA kit procedure. Overall, the highest concentration of copper (1312 mg/kg) induced the most severe damages to plants compared to the 656 mg/kg and 130 mg/kg treatments. Copper at the 1312 mg/kg concentration induced an upregulation of the MRP4 gene in roots and needles. The 656 mg/kg and 130 mg /kg of copper induced no significant difference in MRP4 expression compared to the controls. An upregulation of the RAN1 gene was observed only in roots treated with the 1312 mg/kg. This suggests that the amounts of bioavailable copper in mining sites in Northern Ontario cannot induce changes in MRP4 and RAN1 expressions. Differential expressions of RAN1 and MRP4 genes were observed in roots of copper-resistant and copper-susceptible genotypes. An increase of MRP4 gene expression also observed in the resistant and susceptible genotypes compared to the water control. Copper did not induce changes in the level of global cytosine methylation. However, the potassium ions used as control induced a hypomethylation of DNA at the 1312 mg/kg concentration. [ABSTRACT FROM AUTHOR]

Light availability drives vertical canopy gradients in photosynthetic functioning and carbon (C) balance, yet patterns of variability in these gradients remain unclear. We measured light availability, photosynthetic CO2 and light response curves, foliar C, nitrogen (N) and pigment concentrations, and the photochemical reflectance index (PRI) on upper and lower canopy needles of white spruce trees (Picea glauca) at the species' northern and southern range extremes. We combined our photosynthetic data with previously published respiratory data to compare and contrast canopy C balance between latitudinal extremes. We found steep canopy gradients in irradiance, photosynthesis and leaf traits at the southern range limit, but a lack of variation across canopy positions at the northern range limit. Thus, unlike many tree species from tropical to mid‐latitude forests, high latitude trees may not require vertical gradients of metabolic activity to optimize photosynthetic C gain. Consequently, accounting for self‐shading is less critical for predicting gross primary productivity at northern relative to southern latitudes. Northern trees also had a significantly smaller net positive leaf C balance than southern trees suggesting that, regardless of canopy position, low photosynthetic rates coupled with high respiratory costs may ultimately constrain the northern range limit of this widely distributed boreal species. Summary statement: Canopy gradients in photosynthetic capacity of white spruce diminish at high compared to low latitudes. Low carbon balance in high latitude trees may determine the extent of northern treeline. [ABSTRACT FROM AUTHOR]

In recent decades, forest nurseries in eastern Canada have been faced with periods of mild winter weather, delayed snowfall, and low seedling protective snow cover combined with winter rains instead of snowfall. These extreme conditions have resulted in the loss of millions of seedlings, in particular those that overwinter outdoors, probably due to their winter dehardening. The main objective of this study is to simulate different periods of warm weather at the beginning and end of winter and evaluate their effects on the dehardening and growth of Picea mariana and Picea glauca seedlings in response to different freezing temperatures. Three warming treatments were simulated (control, 1 day, and 3 days of warming at 10 °C) followed by three freezing temperatures (−4, −12, and −20 °C). In winter, regardless of the warming treatment, the seedlings of the two species tolerated the different freezing temperatures without any apparent damage. However, at the end of winter and in the absence of snow cover, the seedlings did not show frost tolerance at −20 °C. On the other hand, the seedlings showed normal growth after undergoing frosts at −4 °C and −12 °C, similar to that observed for control seedlings. Different cultural practices and protection strategies are proposed to improve frost tolerance and reduce the winter loss of seedlings. [ABSTRACT FROM AUTHOR]

Phosphogypsum (PG) is a by-product of phosphorus fertilizer production that is stored in large piles (stacks). Typically, PG stack reclamation focuses on topsoil caps vegetated with grass which has limited economic return. Therefore, a study was conducted using the short-rotational tree species (Picea glauca, Populus balsamifera) to determine their potential in PG stack reclamation. Five soil amendments (compost, compost with mycorrhizal fungi, manure pellets, black earth, mycorrhizal fungi) with a control (no amendment on PG) were used in a field experiment to determine whether they would enhance the growth of the planted species. After two years, amendments had little effect on the height, diameter or biomass of either species. P. balsamifera on PG (control) was significantly healthier than in amended soils. The healthiest and most successful trees for both species were found in the control treatment. Organic amendments (compost, manure) had significantly higher nitrate and phosphorus than most of the other treatments. Black earth and mycorrhizal fungi did not affect any soil properties relative to the control. This study suggests short-rotational forestry plantations of Populus balsamifera appear to be meeting reclamation objectives for PG stacks, similar to traditional grass covers; however, the built soil profile could limit tree success in future. [ABSTRACT FROM AUTHOR]

Climate change is increasing the frequency and intensity of drought events in many boreal forests. Trees are sessile organisms with a long generation time, which makes them vulnerable to fast climate change and hinders fast adaptations. Therefore, it is important to know how forests cope with drought stress and to explore the genetic basis of these reactions. We investigated three natural populations of white spruce (Picea glauca) in Alaska, located at one drought-limited and two cold-limited treelines with a paired plot design of one forest and one treeline plot. We obtained individual increment cores from 458 trees and climate data to assess dendrophenotypes, in particular the growth reaction to drought stress. To explore the genetic basis of these dendrophenotypes, we genotyped the individual trees at 3000 single nucleotide polymorphisms in candidate genes and performed genotype-phenotype association analysis using linear mixed models and Bayesian sparse linear mixed models. Growth reaction to drought stress differed in contrasting treeline populations. Therefore, the populations are likely to be unevenly affected by climate change. We identified 40 genes associated with dendrophenotypic traits that differed among the treeline populations. Most genes were identified in the drought-limited site, indicating comparatively strong selection pressure of drought-tolerant phenotypes. Contrasting patterns of drought-associated genes among sampled sites and in comparison to Canadian populations in a previous study suggest that drought adaptation acts on a local scale. Our results highlight genes that are associated with wood traits which in turn are critical for the establishment and persistence of future forests under climate change. [ABSTRACT FROM AUTHOR]

Under climate change circumstances, increasing studies have reported the temporal instability of tree growth responses to climate, which poses a major challenge to linearly extrapolating past climate and future growth dynamics using tree‐ring data. Space‐for‐time substitution (SFTS) is a potential solution to this problem that is widely used in the dendrochronology field to project past or future temporal growth response trajectories from contemporary spatial patterns. However, the projected accuracy of the SFTS in the climate effects on tree growth remains uncertain. Here, we empirically test the SFTS method by comparing the effect of spatial and temporal climate variations on climate responses of white spruce (Picea glauca), which has a transcontinental range in North America. We first applied a response surface regression model to capture the variations in growth responses along the spatial climate gradients. The results showed that the relationships between growth and June temperature varied along spatial climate gradients in a predictable way. And their relationships varied mainly along with local temperate condition. Then, the projected correlation coefficients between growth and climate using SFTS were compared against the observed. We found that the growth response changes caused by spatial versus temporal climate variations showed opposite trends. Moreover, the projected correlation coefficients using the SFTS were significantly lower than the observed. This finding suggests that applying the SFTS to project the growth response of white spruce might lead to an overestimation of the degree of tree maladaptation in future climate scenarios. And the overestimation is likely to get weaker from Alaska and Yukon Territory in the west to Quebec in the east. Although this is only a case study of the SFTS method for projecting tree growth response, our findings suggest that direct application of the SFTS method may not be applicable to all regions and all tree species. [ABSTRACT FROM AUTHOR]

Recent climate warming is expected to increase tree growth and productivity, substantially altering ecological function and boundaries in northern ecosystems. Although climate largely determines the range and growth of trees in any biome, variations in microsite conditions can influence growth on a finer scale. Red foxes (Vulpes vulpes) are found in most terrestrial ecosystems and are considered ecosystem engineers through their denning activities. Added soil nutrients from prey remains, feces, and urine could benefit tree growth on dens in subarctic regions by alleviating soil nutrient limitations. We examined growth in white spruce (Picea glauca) trees growing on eight red fox dens and paired control sites at the Arctic treeline. Radial growth was 55% higher for trees on dens than on control sites between 1897 and 2017, despite similarities in tree ages and densities. This enhanced growth was more pronounced in younger trees, consistent with the decrease in growth with age among all trees. By promoting tree growth near the treeline, red foxes may act as hotspots for tree reproduction. Although the impacts on tree growth largely depend on the spatial distribution of dens, predators can create distinct microhabitats across the landscape, leading to increased vegetation productivity, persisting over many decades. [ABSTRACT FROM AUTHOR]

White spruce (Picea glauca) spans a massive range, yet the variability in respiratory physiology and related implications for tree carbon balance at the extremes of this distribution remain as enigmas. Working at both the most northern and southern extents of the distribution range more than 5000 km apart, we measured the short‐term temperature response of dark respiration (R/T) at upper and lower canopy positions. R/T curves were fit to both polynomial and thermodynamic models so that model parameters could be compared among locations, canopy positions, and with previously published data. Respiration measured at 25°C (R25) was 68% lower at the southern location than at the northern location, resulting in a significantly lower intercept in R/T response in temperate trees. Only at the southern location did upper canopy leaves have a steeper temperature response than lower canopy leaves, likely reflecting canopy gradients in light. At the northern range limit respiration is nearly twice that of the average R25 reported in a global leaf respiration database. We predict that without significant thermal acclimation, respiration will increase with projected end‐of‐the‐century warming and will likely constrain the future range limits of this important boreal species. Summary Statement: White spruce (Picea glauca) needle respiration at the northern limit of the species range is three times higher than at the southern range limit (when measured at 25°C). This high carbon cost likely challenges tree survival and contributes to the location of the northern treeline. [ABSTRACT FROM AUTHOR]

Spruce–fir (Picea–Abies) forests of the North American Acadian Forest Region are at risk of disappearing from the northeastern United States and Canada due to climate change. Species distribution models (SDMs) have been used to predict changes in this critical transitional ecosystem in the past, but none have addressed how seasonal patterns of temperature and precipitation interact to influence tree species abundance. Inferences have also been limited by contemporary inventory data that could not fully characterize species ranges because they either, (1) only sampled species occurrence after large‐scale human disturbance and settlement, or (2) did not span critical geopolitical boundaries (e.g., the US–Canadian border) that intersect the focal species' range(s). Here, we built new SDM models to better assess the bioclimatic distribution of four spruce–fir species and to test the importance of seasonal climate interactions. We compiled an extensive database of tree occurrence and abundance from recent (~1955–2012) and historical time periods (1623–1869) to model current species distributions and to predict how these might change under future climate. We found that including historical tree data in our SDMs revealed previously unrecognized suitable habitat along the southern edge of species' contemporary ranges. Random forest models predicted occurrence with high accuracy (area under receiver operator curve >0.98), and the seasonal climate variables that emerged as most important for these cold‐adapted species all included interactions that reflected sensitivity to colder temperatures, and preferences for wet weather concentrated in the winter months. Under moderate climate warming (representative concentration pathway 6.0), the northeastern United States retained additional suitable habitat when historical data were included through 2060 for three of the four species: red spruce (Picea rubens), black spruce (Picea mariana), and balsam fir (Abies balsamea), while white spruce (Picea glauca) habitat contracted into Canada. In contrast, future predictions from models that used contemporary data alone forecast extirpation for all four species from the northeastern United States. Overall, these findings highlight that prediction of species ranges in transitional ecosystems that span geopolitical boundaries and gradients of intense land use are improved when historical data and seasonal climate interactions of both temperature and precipitation variables are incorporated. [ABSTRACT FROM AUTHOR]

Merging robust statistical methods with complex simulation models is a frontier for improving ecological inference and forecasting. However, bringing these tools together is not always straightforward. Matching data with model output, determining starting conditions, and addressing high dimensionality are some of the complexities that arise when attempting to incorporate ecological field data with mechanistic models directly using sophisticated statistical methods. To illustrate these complexities and pragmatic paths forward, we present an analysis using tree‐ring basal area reconstructions in Denali National Park (DNPP) to constrain successional trajectories of two spruce species (Picea mariana and Picea glauca) simulated by a forest gap model, University of Virginia Forest Model Enhanced—UVAFME. Through this process, we provide preliminary ecological inference about the long‐term competitive dynamics between slow‐growing P. mariana and relatively faster‐growing P. glauca. Incorporating tree‐ring data into UVAFME allowed us to estimate a bias correction for stand age with improved parameter estimates. We found that higher parameter values for P. mariana minimum growth under stress and P. glauca maximum growth rate were key to improving simulations of coexistence, agreeing with recent research that faster‐growing P. glauca may outcompete P. mariana under climate change scenarios. The implementation challenges we highlight are a crucial part of the conversation for how to bring models together with data to improve ecological inference and forecasting. [ABSTRACT FROM AUTHOR]

Arctic Treeline is the transition from the boreal forest to the treeless tundra and may be determined by growing season temperatures. The physiological mechanisms involved in determining the relationship between the physical and biological environment and the location of treeline are not fully understood. In Northern Alaska, we studied the relationship between temperature and leaf respiration in 36 white spruce (Picea glauca) trees, sampling both the upper and lower canopy, to test two research hypotheses. The first hypothesis is that upper canopy leaves, which are more directly coupled to the atmosphere, will experience more challenging environmental conditions and thus have higher respiration rates to facilitate metabolic function. The second hypothesis is that saplings [stems that are 5–10cm DBH (diameter at breast height)] will have higher respiration rates than trees (stems ≥10cm DBH) since saplings represent the transition from seedlings growing in the more favorable aerodynamic boundary layer, to trees which are fully coupled to the atmosphere but of sufficient size to persist. Respiration did not change with canopy position, however respiration at 25°C was 42% higher in saplings compared to trees (3.43±0.19 vs. 2.41±0.14μmolm−2 s−1). Furthermore, there were significant differences in the temperature response of respiration, and seedlings reached their maximum respiration rates at 59°C, more than two degrees higher than trees. Our results demonstrate that the respiratory characteristics of white spruce saplings at treeline impose a significant carbon cost that may contribute to their lack of perseverance beyond treeline. In the absence of thermal acclimation, the rate of leaf respiration could increase by 57% by the end of the century, posing further challenges to the ecology of this massive ecotone. [ABSTRACT FROM AUTHOR]

Treeline ecosystems are of great scientific interest to study the effects of limiting environmental conditions on tree growth. However, tree growth is multidimensional, with complex interactions between height and radial growth. In this study, we aimed to disentangle effects of height and climate on xylem anatomy of white spruce [ Picea glauca (Moench) Voss] at three treeline sites in Alaska; i.e., one warm and drought-limited, and two cold, temperature-limited. To analyze general growth differences between trees from different sites, we used data on annual ring width, diameter at breast height (DBH), and tree height. A representative subset of the samples was used to investigate xylem anatomical traits. We then used linear mixed-effects models to estimate the effects of height and climatic variables on our study traits. Our study showed that xylem anatomical traits in white spruce can be directly and indirectly controlled by environmental conditions: hydraulic-related traits seem to be mainly influenced by tree height, especially in the earlywood. Thus, they are indirectly driven by environmental conditions, through the environment's effects on tree height. Traits related to mechanical support show a direct response to environmental conditions, mainly temperature, especially in the latewood. These results highlight the importance of assessing tree growth in a multidimensional way by considering both direct and indirect effects of environmental forcing to better understand the complexity of tree growth responses to the environment. [ABSTRACT FROM AUTHOR]

Conifers are the world's major source of timber and pulpwood and have great economic and ecological value. Currently, little research on the application of CRISPR/Cas9, the commonly used genome-editing tool in angiosperms, has been reported in coniferous species. An efficient CRISPR/Cas9 system based on somatic embryogenesis (SEis) suitable for conifers could benefit both fundamental and applied research in these species. In this study, the SpCas9 gene was optimized based on codon bias in white spruce, and a spruce U6 promoter was cloned and function-validated for use in a conifer specific CRISPR/Cas9 toolbox, i.e., PgCas9/PaU6. With this toolbox, a genome-editing vector was constructed to target the DXS1 gene of white spruce. By Agrobacterium -mediated transformation, the genome-editing vector was then transferred into embryogenic tissue of white spruce. Three resistant embryogenic tissues were obtained and used for regenerating plants via SEis. Albino somatic embryo (SE) plants with mutations in DXS1 were obtained in all of the three events, and the ratios of the homozygous and biallelic mutants in the 18 albino mutants detected were 22.2% in both cases. Green plants with mutations in DXS1 were also produced, and the ratios of the DXS1 mutants to the total green plants were 7.9, 28, and 13.5%, respectively, among the three events. Since 22.7% of the total 44 mutants were edited at both of the target sites 1 and 2, the CRISPR/Cas9 toolbox in this research could be used for multi-sites genome editing. More than 2,000 SE plants were regenerated in vitro after genome editing, and part of them showed differences in plant development. Both chimerism and mosaicism were found in the SE plants of white spruce after genome editing with the CRISPR/Cas9 toolbox. The conifer-specific CRISPR/Cas9 system developed in this research could be valuable in gene function research and trait improvement. [ABSTRACT FROM AUTHOR]

Cryopreservation is one of the key technologies for the mass propagation of conifers via somatic embryogenesis. Cryotolerance and embryogenecity of conifer somatic embryos (SEs) could be affected by different temperature treatments, for which the underlying mechanisms were unknown. In this study, the developing SEs of Picea glauca obtained their cryotolerance with a survival rate of 100% when cultured on maturation medium at either 23°C for 4 weeks or 4°C for 10 weeks. However, only the embryos that underwent 4°C acclimation remained high embryogenicity, i.e., 91.7% based on cryovials or 29.3% on the plant tissue. Analysis of differentially expressed genes (DEGs) revealed that both 23 and 4°C treatments led to drastic changes in the gene expression, i.e., 21,621 and 14,906 genes, respectively, and the general increase in many oligosaccharides and flavonoids, in addition to the content change of proline (1.9- and 2.3-fold at 23 or 4°C) and gallic acid (6,963- and 22,053-fold). There were 249 significantly different metabolites between the samples of 23 and 4°C treatments and the changing trend of the sorbitol, fatty acids, and monosaccharides differed between these samples. During 4°C-acclimation, the metabolites of the arginine biosynthesis pathway increased between 2.4- and 8.1-fold, and the expression of antioxidant genes was up-regulated significantly. At 4°C, the up-regulated genes were for germ-like proteins, instead of seed storage proteins at 23°C. Concentrations of abscisic acid and jasmonic acid increased up to 2- and 1.5-fold, respectively, in the cold-acclimated embryos. After 10 weeks at 4°C, the embryos stayed at pre-cotyledonary stage with 17.1% less DNA methylation and fewer storage substances than those at 23°C for 4 weeks, which developed cotyledons. This research provides new insights into mechanisms underlying the response of SEs to different culture temperatures and benefits method development for germplasm conservation in conifers. [ABSTRACT FROM AUTHOR]

Growing Picea glauca seedlings poses many challenges for forest nursery managers, includ- ing reaching the target height at the end of the first growing season (1 + 0) and the homogeneity and uniformity of seedling growth. To increase growth and morphophysiological quality of white spruce seedlings (1 + 0), emphasis was placed on improving the physicochemistry of the acidic peat substrate by replacing silica with granular calcite as a covering material. The objective of our study is to compare the effects of silica and calcite on the physicochemistry of the peat substrate, as well as on growth and mineral nutrition of white spruce seedlings (1 + 0) under forest nursery conditions. Three treatments were used to cover the cavities of large white spruce seedlings produced in containers: silica (29 g/cavity; control treatment), calcite (24 g/cavity) and calcite + (31 g/cavity). At the end of the first growing season, the two calcite treatments significantly increased the total dry mass (28%), roots (27%) and shoot (29%) dry masses and height (24%) compared to silica treatments. Average calcium concentration and content of calcite treatments were significantly higher than that of silica treatment. [ABSTRACT FROM AUTHOR]

The ranges of black and white spruce are largely sympatric, suggesting both species have similar climate requirements. The two species, however, are highly segregated across the landscape with black spruce most common on nutrient‐poor sites with cold, poorly drained soils and white spruce more common on productive sites with warmer, well‐drained soils. Because site conditions influence tree climate–growth responses, it is difficult to compare white and black spruce climate–growth responses as these responses are confounded by the differences in site conditions in which the two species naturally occur. As the climate warms dramatically in northern latitudes, it is critical to understand how a changing climate and associated changes in permafrost and fire regimes will interact to shape future species composition and ecosystem functioning in the boreal forest. In this study, we examined the climate–growth responses of black and white spruce growing in the same sites. This approach eliminates the confounding factor of site conditions and facilitates our understanding of how these two species respond to climate. We included standardized thaw depth of the active layer in our analysis as a representation of permafrost, which is a key factor delineating these two species' habitat preferences and is actively warming and thawing as the climate warms. Our most important finding was that the climate–growth responses of the two species, but especially white spruce, hinged on the thaw depth of the active layer. Specifically, with increasing June–July temperatures white spruce radial growth increased in areas with deep thaw or no near‐surface permafrost, but strongly decreased when growing in areas with near‐surface permafrost. Black spruce radial growth was less sensitive to June–July temperature than white spruce but had a consistent and more positive response to summer precipitation. These findings point to a primary mechanism potentially driving the positioning of these two tree species within the landscapes of boreal interior Alaska and imply widespread thawing of permafrost may foster expansion of white spruce in this region at the expense of black spruce, but that in a wetter climate, black spruce may gain competitive advantage over white spruce in some landscape positions. [ABSTRACT FROM AUTHOR]

Numerous recent studies have argued that moisture limitation is leading to growth declines and mortality of black spruce (Picea mariana) and white spruce (Picea glauca) in the North American boreal forest. A parallel line of research suggests that increasingly common severe wildfires are altering successional pathways and leading to long‐term replacement of spruce forests with those dominated by paper birch (Betula papyrifera, Betula neoalaskana) and trembling aspen (Populus tremuloides). When both conifers and deciduous trees establish after fire, this biome shift hypothesis implicitly assumes that deciduous species will outcompete the conifers, owing to their more rapid vertical growth and because they might be less sensitive to warm and dry conditions.We established a research site in a white spruce‐paper birch forest on an east‐facing slope in interior Alaska and tested the hypothesis that Alaska paper birch are better adapted to warm and dry conditions than white spruce. Over 6 years (2013–2018), we made hourly measurements of microclimate and xylem sap flux of both species. We also collected increment cores and conducted climate‐growth analyses for both species.During our 6‐year study, growing seasons with low volumetric soil water content (SWC) were those that followed shallow winter snowpacks and had limited summer rainfall, not necessarily those with warm air temperature. Both species were sensitive to moisture limitation. The tree‐ring data revealed significant positive effects of cumulative water year precipitation on radial growth, with a steeper slope for paper birch than for white spruce. Radial growth of both species was also positively related to mean water year air temperature. Sap flux density declined progressively for white spruce over the range of observed SWC and abruptly for paper birch when SWC fell below ~15%.Synthesis. Our results show that, while paper birch might be less sensitive to mild drought than white spruce, it may be more sensitive to severe drought, raising questions about the ability of paper birch to outcompete co‐occurring white spruce in a drier climate. [ABSTRACT FROM AUTHOR]

Conifer forests worldwide are becoming increasingly vulnerable to the effects of climate change. Although the production of phenolic compounds (PCs) has been shown to be modulated by biotic and abiotic stresses, the genetic basis underlying the variation in their constitutive production level remains poorly documented in conifers. We used QTL mapping and RNA-Seq to explore the complex polygenic network underlying the constitutive production of PCs in a white spruce (Picea glauca) full-sib family for 2 years. QTL detection was performed for nine PCs and differentially expressed genes (DEGs) were identified between individuals with high and low PC contents for five PCs exhibiting stable QTLs across time. A total of 17 QTLs were detected for eight metabolites, including one major QTL explaining up to 91.3% of the neolignan-2 variance. The RNA-Seq analysis highlighted 50 DEGs associated with phenylpropanoid biosynthesis, several key transcription factors, and a subset of 137 genes showing opposite expression patterns in individuals with high levels of the flavonoids gallocatechin and taxifolin glucoside. A total of 19 DEGs co-localized with QTLs. Our findings represent a significant step toward resolving the genomic architecture of PC production in spruce and facilitate the functional characterization of genes and transcriptional networks responsible for differences in constitutive production of PCs in conifers. [ABSTRACT FROM AUTHOR]

• Over many decades, red foxes increase nutrient concentrations and affect vegetation diversity on their dens in the boreal woodlands, a nutrient-poor environment. • Denning activity also affects the growth and reproduction of white spruce, a mast-seeding conifer. • White spruce on fox dens are taller and produce more cones in non-mast years than trees off dens. • The effects of denning are subject to the mast-seeding cycle, as cone production in mast years was not elevated on dens. Predators may act as ecosystem engineers by modifying their physical environment through non-trophic interactions. Red foxes (Vulpes vulpes) are ecosystem engineers in the boreal woodlands, a nutrient-poor environment, where their denning activity increases soil nutrient concentrations, affecting the diversity of vegetation around dens. Since dens can persist for many decades and tree growth is enhanced on dens, dens can be used to study the long term impacts of nutrient additions on community and ecosystem processes. We examined the reproductive output of a mast-seeding conifer, white spruce (Picea glauca), on 10 red fox dens and paired control sites at the boreal treeline near Churchill, Manitoba, in July 2019. We estimated cone production in both the current non-mast year (2019) and the previous mast year. The number of cones produced per tree and per hectare was significantly higher on dens than control sites in the non-mast year, but did not differ in the mast year. Higher cone production on dens was partly driven by trees being larger on dens. These results suggest that red fox ecosystem engineering activity affects white spruce reproduction, since increased soil nutrients on dens allow for higher cone production, but the effect of red fox denning is limited by interactions with mast-seeding mechanisms. For example, weather cues may override the effects of denning in mast years, while trees on dens may be able to allocate more resources to reproduction in non-mast years, but store the same amount of resources as trees off dens to produce similarly large cone crops in mast years. Altered resource availability in this nutrient-limited landscape could have additional ecological implications, by affecting the foraging patterns of seed predators and spruce range expansion. [ABSTRACT FROM AUTHOR]

With climate change, the pressure on tree breeding to provide varieties with improved resilience to biotic and abiotic stress is increasing. As such, pest resistance is of high priority but has been neglected in most tree breeding programs, given the complexity of phenotyping for these traits and delays to assess mature trees. In addition, the existing genetic variation of resistance and its relationship with productivity should be better understood for their consideration in multitrait breeding. In this study, we evaluated the prospects for genetic improvement of the levels of acetophenone aglycones (AAs) in white spruce needles, which have been shown to be tightly linked to resistance to spruce budworm. Furthermore, we estimated the accuracy of genomic selection (GS) for these traits, allowing selection at a very early stage to accelerate breeding. A total of 1,516 progeny trees established on five sites and belonging to 136 full‐sib families from a mature breeding population in New Brunswick were measured for height growth and genotyped for 4,148 high‐quality SNPs belonging to as many genes along the white spruce genome. In addition, 598 trees were assessed for levels of AAs piceol and pungenol in needles, and 578 for wood stiffness. GS models were developed with the phenotyped trees and then applied to predict the trait values of unphenotyped trees. AAs were under moderate‐to‐high genetic control (h2: 0.43–0.57) with null or marginally negative genetic correlations with other traits. The prediction accuracy of GS models (GBLUP) for AAs was high (PAAC: 0.63–0.67) and comparable or slightly higher than pedigree‐based (ABLUP) or BayesCπ models. We show that AA traits can be improved and that GS speeds up the selection of improved trees for insect resistance and for growth and wood quality traits. Various selection strategies were tested to optimize multitrait gains. [ABSTRACT FROM AUTHOR]

Tree growth at northern treelines is generally temperature‐limited due to cold and short growing seasons. However, temperature‐induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree‐ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell‐wall thickness, cell number) and TRW were correlated with the drought‐sensitive standardized precipitation–evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925–1946), cool/wet (1947–1976) and again warm/dry (1977–1998) climate regimes. Xylem anatomical traits revealed water‐limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture‐driven shift in growth‐limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture‐driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity. [ABSTRACT FROM AUTHOR]

Summary: Laimaphelenchus pannocaudus, isolated in Ningbo port from Picea gluaca, is redescribed based upon characteristic morphological details elucidated by light and scanning electron microscopy photographs. The recovered population is characterised by 907 (771-1024) μ m long females with 11.2 (8.9-12.2) μ m long stylet, four incisures in the lateral field, excretory pore located at the same level as, or slightly anterior to the nerve ring, vulva lacking a flap and located at 68.7 (63.9-73.4)% of the body, elongate post-vulval uterine sac 78.5 (55.6-101.1) μ m long, and tail 40.6 (33.4-45.6) μ m long. Males are common and characterised by 16.5 (15.7-17.8) μ m long spicules (chord) with their condylus slightly recurved, squared to rounded in shape with a blunt rounded tip and rostrum triangular with a bluntly pointed tip, and six visible caudal papillae. The female tail is ventrally curved, conoid and bears a stalk-like terminus with 4-8 pedunculate projections. Phylogenetic analyses using partial 18S and 28S rDNA D2-D3 data revealed that L. pannocaudus formed a sister relationship with L. suberensis in both phylogenies. [ABSTRACT FROM AUTHOR]

Intensive plantation forestry is a potent strategy for forest managers to increase wood production on a smaller forest land acreage, especially with the use of genetically improved reforestation stock. The main drawback with conventional conifer improvement is the several decades it takes before stock deployment, which is particularly acute in the context of climate change and evolving wood markets. Use of genomic selection allows to drastically shorten breeding cycles, resulting in more flexibility and potentially increasing benefits. This study compares the financial performance of five white spruce (Picea glauca) breeding and deployment scenarios, from conventional breeding to genomic selection in conjunction with top-grafting or the use of somatic embryogenesis, in the context of plantations established by the Quebec government on public lands with banned herbicide use. We estimated the land expectation value (LEV) for the five scenarios applied to eight site productivity indices, and considered costs and revenues from breeding, plantation establishment, silviculture, and harvest. LEVs at 4% discount rate were positive for all scenarios on all site indices, and varied from $197 to $2015 ha−1 assuming mechanical brushing of the plantations. The scenarios integrating genomic selection resulted in the highest LEVs, which increased with site index. We also conducted sensitivity analyses with 3% and 5% discount rates, with a range of costs and benefits, and with herbicide control of competing vegetation. These results should help orientate public investment decisions regarding the integration of genomic selection at the operational level in tree breeding and reforestation programs on public lands. [ABSTRACT FROM AUTHOR]

The spruce budworm, Choristoneura fumiferana, is presumed to be panmictic across vast regions of North America. We examined the extent of panmixia by genotyping 3,650 single nucleotide polymorphism (SNP) loci in 1975 individuals from 128 collections across the continent. We found three spatially structured subpopulations: Western (Alaska, Yukon), Central (southeastern Yukon to the Manitoba–Ontario border), and Eastern (Manitoba–Ontario border to the Atlantic). Additionally, the most diagnostic genetic differentiation between the Central and Eastern subpopulations was chromosomally restricted to a single block of SNPs that may constitute an island of differentiation within the species. Geographic differentiation in the spruce budworm parallels that of its principal larval host, white spruce (Picea glauca), providing evidence that spruce budworm and spruce trees survived in the Beringian refugium through the Last Glacial Maximum and that at least two isolated spruce budworm populations diverged with spruce/fir south of the ice sheets. Gene flow in the spruce budworm may also be affected by mountains in western North America, habitat isolation in West Virginia, regional adaptations, factors related to dispersal, and proximity of other species in the spruce budworm species complex. The central and eastern geographic regions contain individuals that assign to Eastern and Central subpopulations, respectively, indicating that these barriers are not complete. Our discovery of previously undetected geographic and genomic structure in the spruce budworm suggests that further population modelling of this ecologically important insect should consider regional differentiation, potentially co‐adapted blocks of genes, and gene flow between subpopulations. [ABSTRACT FROM AUTHOR]

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Identifying the main drivers of tree height growth on reclaimed oil sands sites of northern Alberta can provide useful information on what can be done to shorten the recovery time of these disturbed sites. The effect of water availability, foliar nutrient concentrations, competition, and soil chemical properties on young trembling aspen and white spruce height across two soil types (peat-mineral mix and forest floor-mineral mix) and two fertilizer levels (200 kg NPK ha−1 and no fertilizer) was examined. For comparison, we also studied a natural site that burned the same year the reclaimed site was established. On the reclaimed site, peat-mineral soil had the greatest tree height for both species, though not by a significant margin in the case of white spruce. The fertilizer treatment had no apparent effect on tree height on the forest floor-mineral soil and a negative effect on the peat-mineral soil. Fertilization increased vegetation cover on both soil types, which, when combined with the negative or lack of durable effect of fertilization on tree height, suggest that the increase in competition for resources could have had a stronger effect on tree height than the increase in immediately available nutrients following fertilization. The majority of tree height growth drivers found were related to nutrient availability and, as far as we can tell, did not significantly differ between reclamation soil types. Aspen sucker height on the burned site was not strongly correlated to the measured variables, suggesting that suckers do not yet fully rely on their environment for height growth. [ABSTRACT FROM AUTHOR]

A cost‐effective climate change adaptation strategy for the forestry sector is to move seed sources to more northern and higher elevation planting sites as part of ongoing reforestation programs. This is meant to match locally adapted populations with anticipated environments, but adaptive traits do not always show population differences suitable to mitigate climate change impacts. For white spruce, drought tolerance is a critical adaptive trait to prevent mortality and productivity losses. Here, we use a 40‐year‐old provenance experiment that has been exposed to severe drought periods in 1999 and 2002 to retrospectively investigate drought response and the adaptive capacity of white spruce populations across their boreal range. Relying on dendrochronological analysis under experimentally controlled environments, we evaluate population differences in resistance, resilience, and recovery to these extreme events. Results showed evidence for population differentiation in resistance and recovery parameters, but provenances conformed to approximately the same growth rates under drought conditions and had similar resilience metrics. The lack of populations with better growth rates under drought conditions is contrary to expectations for a wide‐ranging species with distinct regional climates. Populations from the wettest environments in the northeastern boreal were surprisingly drought‐tolerant, suggesting that these populations would readily resist water deficits projected for the 2080s, and supporting the view that northeastern Canada will provide a refugium for boreal species under climate change. The findings also suggest that white spruce is sensitive to growth reductions under climate change in the western boreal. The study highlights that population differentiation in adaptive capacity is species‐ and trait‐specific, and we provide a counterexample for drought tolerance traits, where assisted migration prescriptions may be ineffective to mitigate climate change impacts. For resource managers and policy makers, we provide maps where planning for widespread declines of boreal white spruce forests may be unavoidable. [ABSTRACT FROM AUTHOR]

Under a changing climate, tree improvement programs should revise breeding zones to improve climate change adaptation and program efficiency once information about adaptive capacity of breeding populations and genotype × environment (G × E) interaction becomes available. In this study, we evaluated the adaptation and growth, as well as G × E interactions, shown by white spruce (Picea glauca (Moench) Voss) open-pollinated (OP) families across two adjacent breeding zones (i.e., the BZ3300 and BZ4300, respectively) in northwestern Ontario using progeny test data, including 80 OP families (40 from each) that were cross-tested between the breeding zones. Results showed that the OP families from the southerly BZ4300 breeding zone averaged higher survival and significantly greater mean tree height growth than the local OP families in the more northern BZ3300 breeding zone at tree age of 14 years, with no notable difference in frost damage. Estimates of type-B genetic correlation from pairwise tests or combined-test analyses were almost all greater than 0.7, suggesting weak G × E interactions. Greater variation among families and higher heritability estimates in tree height were observed for families from the BZ3300 breeding zone or when families from the two breeding zones were combined. The weak G × E interactions, in addition to better survival and growth of the southerly families in the more northern breeding zone, suggested benefits from deploying seed from current seed orchard in the BZ4300 breeding zone to the BZ3300 breeding zone and merging the two breeding zones when starting the second generation breeding to increase tree improvement efficiency and likely higher genetic gain under future climate. [ABSTRACT FROM AUTHOR]

The negative growth response of North American boreal forest trees to warm summers is well documented and the constraint of competition on tree growth widely reported, but the potential interaction between climate and competition in the boreal forest is not well studied. Because competition may amplify or mute tree climate‐growth responses, understanding the role current forest structure plays in tree growth responses to climate is critical in assessing and managing future forest productivity in a warming climate. Using white spruce tree ring and carbon isotope data from a long‐term vegetation monitoring program in Denali National Park and Preserve, we investigated the hypotheses that (a) competition and site moisture characteristics mediate white spruce radial growth response to climate and (b) moisture limitation is the mechanism for reduced growth. We further examined the impact of large reproductive events (mast years) on white spruce radial growth and stomatal regulation. We found that competition and site moisture characteristics mediated white spruce climate‐growth response. The negative radial growth response to warm and dry early‐ to mid‐summer and dry late summer conditions intensified in high competition stands and in areas receiving high potential solar radiation. Discrimination against 13C was reduced in warm, dry summers and further diminished on south‐facing hillslopes and in high competition stands, but was unaffected by climate in open floodplain stands, supporting the hypothesis that competition for moisture limits growth. Finally, during mast years, we found a shift in current year's carbon resources from radial growth to reproduction, reduced 13C discrimination, and increased intrinsic water‐use efficiency. Our findings highlight the importance of temporally variable and confounded factors, such as forest structure and climate, on the observed climate‐growth response of white spruce. Thus, white spruce growth trends and productivity in a warming climate will likely depend on landscape position and current forest structure. Using tree ring and carbon isotopes data from Denali National Park and Preserve, we investigated whether competition and site moisture characteristics mediate white spruce radial growth response to climate. We found competition and solar radiation mediate the influence of climate on the radial growth. Our carbon isotope analysis suggests the mechanism behind these modified climate‐growth responses is increased competition for moisture in high basal area stands and dry sites. Overall, our results suggest dynamic growth responses to future climate change that are dependent on landscape position and stand competition and likely to result in feedbacks on forest structure. [ABSTRACT FROM AUTHOR]

White spruce [ Picea glauca (Moench) Voss] is a commercially valuable boreal tree that has been known for its ability to colonize deglaciated rock tailings. Over the last decade, there has been an increasing interest in using this species for the revegetation and successful restoration of abandoned mine spoils. Herein, we conducted a glasshouse experiment to screen mycorrhizal fungi and rhizobacteria capable of improving the health and growth of white spruce seedlings growing directly on waste rocks (WRs) or fine tailings (FTs) from the Sigma-Lamaque gold mine located in the Canadian Abitibi region. After 32 weeks, measurements of health, growth, and mycorrhizal colonization variables of seedlings were performed. Overall, symbionts isolated from roots of healthy white spruce seedlings growing on the mining site, especially Cadophora finlandia Cad. fin. MBN0213 GenBank No. KC840625 and Pseudomonas putida MBN0213 GenBank No. AY391278, were more efficient in enhancing seedling health and growth than allochthonous species and constitute promising microbial symbionts. In general, mycorrhizae promoted plant health and belowground development, while rhizobacteria enhanced aboveground plant biomass. The observed beneficial effects were substrate-, strain-, and/or strains combination-specific. Therefore, preliminary experiments in control conditions such as the one described here can be part of an efficient and integrated strategy to select ecologically well-adapted symbiotic microorganisms, critical for the success of a long-term revegetation program. [ABSTRACT FROM AUTHOR]

In the context of a phytorestoration project, the purpose of this study was to assess the respective contribution to the nutritional status of Picea glauca seedlings of ectomycorrhizae and rhizobacteria native or not to the Sigma-Lamaque gold mine wastes in northern Quebec, Canada. In a glasshouse experiment, inoculated plants were grown for 32 weeks on coarse waste rocks or fine tailings obtained from the mining site. The survival, health, growth, and nutritional status of plants were better on coarse waste rocks than on fine tailings. Fe and Ca were especially found at high levels in plant tissues but at much lower concentrations on waste rocks. Interestingly, inoculation of microsymbionts had only minimal effects on N, P, K, and Mg plant status that were indeed close or within the concentration range encountered in healthy seedlings. However, both fungal and bacterial treatments improved Fe and Ca concentrations in plant tissues. Fe concentration in the foliage of plants inoculated with the fungi Tricholoma scalpturatum Tri. scalp. MBN0213 GenBank #KC840613 and Cadophora finlandia Cad. fin. MBN0213 GenBank #KC840625 was reduced by >50%. Both fungi were isolated from the mining site. The rhizobacteria, Azotobacter chroococcum , also improved plant Fe level in some cases. Regarding Ca nutritional status, the native bacterial strain Pseudomonas putida MBN0213 GenBank #AY391278 was the only symbiont that reduced foliar content by up to 23%. Ca concentration was negatively correlated with the fungal mycorrhization rate of seedling roots. This relation was especially strong (r = -0.66, p -value ≤ 0.0001) in the case of C. finlandia. Also, a similar relationship existed with root Fe concentration (r = -0.44, p -value ≤ 0.0001). In fact, results showed that seedling performance was more correlated with elevated Ca and Fe concentration in planta than with nutrient deficiency. Also, native microsymbionts were capable of regulating seedling nutrition in the poor substrate of the Sigma-Lamaque gold mine tailings. [ABSTRACT FROM AUTHOR]

Abstract: The boreal biome represents approximately one third of the world's forested area and plays an important role in global biogeochemical and energy cycles. Numerous studies in boreal Alaska have concluded that growth of black and white spruce is declining as a result of temperature‐induced drought stress. The combined evidence of declining spruce growth and changes in the fire regime that favor establishment of deciduous tree species has led some investigators to suggest the region may be transitioning from dominance by spruce to dominance by deciduous forests and/or grasslands. Although spruce growth trends have been extensively investigated, few studies have evaluated long‐term radial growth trends of the dominant deciduous species (Alaska paper birch and trembling aspen) and their sensitivity to moisture availability. We used a large and spatially extensive sample of tree cores from interior Alaska to compare long‐term growth trends among contrasting tree species (white and black spruce vs. birch and aspen). All species showed a growth peak in the mid‐1940s, although growth following the peak varied strongly across species. Following an initial decline from the peak, growth of white spruce showed little evidence of a trend, while black spruce and birch growth showed slight growth declines from ~1970 to present. Aspen growth was much more variable than the other species and showed a steep decline from ~1970 to present. Growth of birch, black and white spruce was sensitive to moisture availability throughout most of the tree‐ring chronologies, as evidenced by negative correlations with air temperature and positive correlations with precipitation. However, a positive correlation between previous July precipitation and aspen growth disappeared in recent decades, corresponding with a rise in the population of the aspen leaf miner (Phyllocnistis populiella), an herbivorous moth, which may have driven growth to a level not seen since the early 20th century. Our results provide important historical context for recent growth and raise questions regarding competitive interactions among the dominant tree species and exchanges of carbon and energy in the warming climate of interior Alaska. [ABSTRACT FROM AUTHOR]

Significant treeline advance can only occur with successful establishment, growth, and survival of new seedlings. Several studies have examined microsite factors at single locations to explain the presence/absence of seedlings at treeline. We conducted a much larger observational study and included multiple factors to determine (1) which variable(s) was/were most important, (2) whether their importance differed between aspects, and (3) whether the same variables explained variation in seedling growth and damage. We analyzed five biophysical and six shrub variables along four forest-tundra ecotones in southwest Yukon at 640 points. The model that best explained seedling occurrence was similar between north- and southfacing slopes. Of all variables, seedling occurrence was best explained by the proximity, height, and upslope orientation of shrubs (relative to the seedling). The data indicate an optimal range of shrub cover, which differed with aspect. On north-facing slopes, seedlings occurred most often when shrub cover exceeded 13%, while on south-facing slopes seedlings occurred most when shrub cover was between 9% and 72%.We also found that with the exception of shrub-related factors, very few biophysical variables explained size and growth characteristics and damage of tree seedlings, suggesting that the relative strength and importance of variables change depending on the life stage and size of the tree seedling. Collectively, our results demonstrate a non-random distribution of seedlings in the forest-tundra ecotone, suggesting that as shrub distributions change with climate change, colonization sites for seedlings will also be influenced. [ABSTRACT FROM AUTHOR]

Since 2001, climatic conditions have been notably drier than normal across large areas of the western Canadian interior, leading to widespread impacts on the forests of this region. This poses a major concern for the future, given climate change projections for continued warming and drying. We conducted tree-ring analysis in 75 pure stands of white spruce ( Picea glauca) across Alberta and west-central Saskatchewan to examine the effects of recent climatic drying on the growth of this important boreal tree species. Allometric equations were used to calculate annual growth in aboveground tree biomass ( GBM) from ring width measurements. Results showed an increasing trend in GBM from the 1960s to the 1990s, followed by a sharp decline during the severe drought of 2001-2002. Of the 75 stands, only 18 recovered sufficiently to cause an increase in mean GBM from the predrought decade of 1991-2000 to the subsequent decade of 2001-2010. The remaining 57 stands exhibited a decline in mean GBM between these decades. Climatic drying was a major cause of the growth decline, as shown by the significant stand-level relationship between percentage change in decadal mean GBM and the change in decadal mean values of a climate moisture index from 1991-2000 to 2001-2010. Subsequent analyses of boreal stands sampled across Alberta during 2015 revealed that white spruce growth had declined even further as drought conditions intensified during 2014-2015. Overall, there was a 38% decrease in mean GBM between 1997 and 2015, but surprisingly, the percentage decrease was not significantly different for young, productive stands compared with older, less productive stands. Thus, stand ageing cannot explain the observed decline in white spruce growth during the past quarter century, suggesting that these forests are at risk if the trend towards more frequent, severe drought continues in the region. [ABSTRACT FROM AUTHOR]

Regional warming has led to increased productivity near the boreal forest margin in Alaska. To date, the effects of warming on seedling recruitment have received little attention, in spite of forecasted forest expansion. Here, we used stand structure and environmental data from 95 white spruce ( Picea glauca) plots sampled across a longitudinal gradient in southwest Alaska to explore factors influencing spruce establishment and recruitment near western treeline. We used total counts of live seedlings, saplings, and trees, representing five life stages, to evaluate whether geospatial, climate, and measured plot covariates predicted abundance, using current abundance distributions as a surrogate for climate conditions in the past. We used generalized linear models to test the null hypothesis that conditions favorable for recruitment were similar along the environmental gradient represented by longitude, by exploring relationships between per-plot counts of each life stage and the covariates hypothesized to affect abundance. We also examined the relationship between growing degree days ( GDD) and seedling establishment over a period of three decades using tree-ring chronologies obtained from cores taken at a subset of our sites ( n = 30). Our results indicated that seedling, sapling, and tree abundance were positively correlated with temperature across the study area. The response to longitude was mixed, with earlier life stages (seedlings, saplings) most abundant at the western end of the gradient, and later life stages (trees) most abundant to the east. The differential relationship between longitude and life-stage abundance suggests a moving front of white spruce establishment through time, driven by changes in environmental conditions near the species' western range limit. Likewise, we found a positive relationship between periods of seedling establishment and GDD, suggesting that longer summers and/or greater heat accumulation might enhance establishment, consistent with the positive relationship we found between life-stage abundance and temperature. [ABSTRACT FROM AUTHOR]

A wide range of Sitka spruce breeding material ( Picea sitchensis (Bong.) Carr.) was assessed for its ability to respond to somatic embryogenesis (SE) and cryo-preservation methods, with 765 cell lines being established from seeds taken from the UK's breeding population for testing with the relevant methodologies. Cell lines were most readily established from immature seeds (63%) on a proliferation medium originally developed for white spruce ( P. glauca) with agar as the gelling agent. However, it was also found that 48% of the zygotic embryos taken from freshly harvested mature seed were also able to establish cell lines on the same medium, as were 20% of seeds that had been stored at −20 °C for up to 8 years. Most of the cell lines tested (57%) produced significant numbers of somatic embryos, but the rate at which they could be germinated into plants capable of surviving the transfer to nursery conditions varied considerably from 67 to 8% between years for reasons that were not clear. The ability of the cell lines to survive cryo-preservation was also assessed, and out of 199 cell lines tested, 45 cell lines from 21 breeding families were successfully established in a cryo-storage facility. The best rates of recovery were obtained when Phytagel was used in the medium for growing the samples before and after they were cryo-preserved. [ABSTRACT FROM AUTHOR]

The effectiveness of nitrogen (N) fertilization in improving tree growth in saline forest soils is poorly understood. Hence, in this study, the interactive effect of N fertilization and salt stress on seedling physiology and growth was examined to improve our understanding of possible mechanisms for fertilization to mitigate salt stress. We compared physiological characteristics, growth, and foliar and root nutrient concentrations in lodgepole pine ( Pinus contorta ) and white spruce ( Picea glauca ) seedlings planted in a peat-mineral soil mix during a three months study with N fertilization (0 and 300 mg N seedling −1 ) and salinity (0, 50, 100, 150 mmol L −1 NaCl) in a greenhouse experiment. Nitrogen fertilization significantly increased net photosynthetic rates ( P n ) and N concentrations in needles and roots, resulting in improved seedling height and root collar diameter growth and total seedling biomass for both species when seedlings were subjected to low salinity (50 mmol L −1 NaCl), while it did not help tree growth under high salinity (150 mmol L −1 NaCl). The physiological responses to combined salt stress and N fertilization did not differ between the two species; however, fertilization improved the growth of pine seedlings more than that of spruce seedlings under different salinity levels. The results suggest that the effectiveness of N fertilization in mitigating the salinity effect on seedling growth was species specific. Our study demonstrates that N fertilization improves tree growth through enhanced photosynthesis and N nutrition under low salinity and provides valuable insights into strategies to manage N nutrition in saline boreal forest soils. [ABSTRACT FROM AUTHOR]

Post-harvest regeneration failure of white spruce ( Picea glauca Moench [Voss]), has led to concerns of “de-coniferization” on productive site in the Alaskan boreal forest. Forest management in the region sought historically to increase spruce composition after harvest through silvicultural practices such as site preparation and assisted regeneration; however, successful reforestation requires the effects of these practices to persist over time and control non-target tree species. In order to identify the enduring effects of silvicultural regeneration practices, we sampled a large (26.7 ha) white spruce regeneration trial established immediately following a stand-replacing wildfire and subsequent salvage harvest in a productive upland forest. The original regeneration treatments followed a split-split plot experimental design on two landform types (LF), four ground scarification treatments (GST) plus a non-scarified control, and five artificial white spruce regeneration treatments (WSRT) plus a natural seedfall control (Densmore et al., 1999). Here we analyze the total biomass, stand density, and basal area for all tree species within each of the regeneration treatments 28 years post-establishment, and calculate seed dispersal distances. Our results show that compared to natural seedfall control plots, white spruce basal area was six times greater in planted seedling plots, and white spruce stem density (dbh ≥ 1.0 cm) was nearly three times greater in broadcast seeding plots. White spruce stem density from natural seedfall averaged 944 stems ha −1 , but was dependent on both topographic position and distance to wind-dispersed seed sources. Our results also indicate that GST had few significant effects on white spruce basal area or stem density. However, scarification nearly doubled Alaska birch ( Betula neoalaskana Sarg.) stem density and basal area compared to non-scarified control plots. Planted white spruce plots supported 19% less birch basal area, except in the most intensive scarification treatments in which birch basal area did not differ. Intensive scarification reduced quaking aspen ( Populus tremuloides Michx.) basal area by half on slope plots. Our results demonstrate that early regeneration practices profoundly influence stand development beyond the stem initiation stage, but pre-fire stand type, post-fire configuration of unburned seed sources, and topographical variation play a mediating role in determining species assemblages and competitive relationships. A fire-killed stand must be considered within its ecological and landscape context to determine the probable success of a management action such as salvage and tree regeneration. [ABSTRACT FROM AUTHOR]

This paper analyzes sapwood variability and allometry within individuals of Populus tremuloides, Pinus contorta, Pinus banksiana, Picea mariana, and Picea glauca. Outside bark diameter at breast height (DBH) and sapwood depth (sd) in four cardinal directions were measured in individuals in stands in Alberta and Saskatchewan, Canada. The microscopical analysis of wood anatomy was used to measure sd, and the error associated with the measures was observed. Sapwood allometry analyses examined the influence of DBH on sd and on sapwood area (SA). All species were observed to have varying sapwood depths around the trunk with statistical analyses showing that Pinus banksiana has a well defined preference to grow thicker in the North-East side. The largest sd values were observed for the Populus tremuloides set. Unlike Populus tremuloides and Picea glauca, for the species Pinus contorta, Pinus banksiana, and Picea mariana, incremental growth in DBH does not directly drive sapwood growth in any direction. For these three species, SA increases only because of increases in DBH as sd remains nearly constant. These results show that sapwood depth and sapwood area seem to behave differently in each studied species and are not always proportional to the tree size as is normally assumed. [ABSTRACT FROM AUTHOR]

While DNA methylation carries genetic signals and is instrumental in the evolution of organismal complexity, small RNAs (sRNAs), ~18-24 ribonucleotide (nt) sequences, are crucial mediators of methylation as well as gene silencing. However, scant study deals with sRNA evolution via featuring their expression dynamics coupled with species of different evolutionary time. Here we report an atlas of sRNAs and microRNAs (miRNAs, single-stranded sRNAs) produced over time at seed-set of two major spermatophytes represented by populations of Picea glauca and Arabidopsis thaliana with different seed-set duration. We applied diverse profiling methods to examine sRNA and miRNA features, including size distribution, sequence conservation and reproduction-specific regulation, as well as to predict their putative targets. The top 27 most abundant miRNAs were highly overlapped between the two species (e.g., miR166,-319 and-396), but in P. glauca, they were less abundant and significantly less correlated with seed-set phases. The most abundant sRNAs in libraries were deeply conserved miRNAs in the plant kingdom for Arabidopsis but long sRNAs (24-nt) for P. glauca. We also found significant difference in normalized expression between populations for population-specific sRNAs but not for lineage-specific ones. Moreover, lineage-specific sRNAs were enriched in the 21-nt size class. This pattern is consistent in both species and alludes to a specific type of sRNAs (e.g., miRNA, tasiRNA) being selected for. In addition, we deemed 24 and 9 sRNAs in P. glauca and Arabidopsis, respectively, as sRNA candidates targeting known adaptive genes. Temperature had significant influence on selected gene and miRNA expression at seed development in both species. This study increases our integrated understanding of sRNA evolution and its potential link to genomic architecture (e.g., sRNA derivation from genome and sRNA-mediated genomic events) and organismal complexity (e.g., association between different sRNA expression and their functionality). [ABSTRACT FROM AUTHOR]

Early establishment of seedlings in reclaimed oil sand areas is often limited by low nutrient and water availability due to factors such as strong understory vegetation competition. Management practices such as nursery fertilization and field weed control could help early establishment of planted seedlings and reclamation success. We investigated the effect of nursery nutrient loading and field weed control on the growth, nitrogen (N) retranslocation within seedling components, and plant N uptake from the soil for white spruce ( Picea glauca [Moench] Voss) seedlings planted on a highly competitive reclaimed oil sands site for two years. Exponential fertilization during nursery production increased the root biomass but not the nutrient reserve in the seedling. In the field experiment, on average across the treatments, 78 and 49% of the total N demand of new tissue growth in the first and second year were met by N retranslocation, respectively. Though exponential fertilization did not affect N retranslocation, it increased the percent height and root collar diameter growth. Weed control increased not only the growth of seedlings by increasing soil N availability, but also N retranslocation within the seedlings in the second year after outplanting. We conclude that vegetation management by weed control is feasible in improving the early growth of white spruce seedlings planted on reclaimed soils and facilitate tree establishment in the oil sands region. Optimization of the nursery exponential N fertilization regime for white spruce may further help with early revegetation of reclaimed oil sands sites. [ABSTRACT FROM AUTHOR]

Aim A critical concern for boreal ecosystems centres on broad-scale responses to warming, i.e. declining growth and mortality, or enhanced growth and greater productivity. However, few studies have synthesized tree growth along biogeographic gradients to address this issue. This study develops a broader understanding of how growth of a dominant conifer has responded to recent warming near the western forest margin of Alaska. Location Alaska, United States. Methods Thirty Picea glauca sites in south-west Alaska (1216 trees ≥ 4 cm dbh) were evaluated for growth differences from the southern boreal forest to the western forest-tundra margin across low-elevation forests and woodlands, and altitudinal tree line. Regional climate records were used to evaluate (1) whether tree growth near western tree line showed greater sensitivity to temperature and/or precipitation than southern boreal sites, (2) if the climate-growth response varied through time, across ecoregions and ecosystems, with spruce beetle disturbance, and by tree age, and (3) if there was a temperature threshold that limited growth. Results Positive growth trends since the 1980s in many open stands were consistent with the predicted expansion of western and altitudinal tree line. However, growth levelled off with temperatures ≥ 13°C at all but altitudinal tree line. An increasingly positive effect of precipitation on growth occurred after 1985, particularly at boreal tree line and closed forest sites. Closed-canopy forests showed lower rates of growth, greater spruce beetle activity and less potential for resiliency to warming than other ecosystem types. Main Conclusions Warming has led to markedly different growth responses according to ecosystem type and ecoregion near the forest-tundra margin. Altitudinal tree line sites showed consistently positive growth with warmer temperatures in recent decades, whereas low-elevation forests had reduced growth and greater beetle mortality. Strong positive correlations between growth and summer-fall precipitation since the mid-1980s suggest that precipitation will become an increasingly important factor with further warming. [ABSTRACT FROM AUTHOR]

Premise of research: Understanding the relative contribution of environmental cues (photoperiod and temperature) as determinants of the transition from active growth to dormancy is important for better predictions of tree responses to anticipated climate change. The aim of this study was to identify the anatomical and structural changes that occur in the cambium of white spruce during the transition from active growth to dormancy triggered by short-day signals. Methodology: Anatomical features at the structural and ultrastructural levels were compared between actively and nonactively growing cambium to better understand how white spruce responds to short photoperiod. Pivotal results: The characteristics observed in the dormant cambium revealed that the organelles are densely and uniformly distributed throughout the cytoplasm. The abundance of small vacuoles, smooth endoplasmic reticulum, and starch granules were the main histological features characterizing the dormant state. In contrast to the dormant cambium cells, the active cells were characterized by abundant ribosomes, golgi bodies, and rough endoplasmic reticulum. Active cambium had cells with smaller radial dimensions and thinner cell walls than those in dormant cambium. Amyloplast-type plastids were also frequently seen and appeared to be empty. Cytoplasm in the active cambium was localized at the periphery of cambial cells and occupied by few large vacuoles in contrast to the numerous small vacuoles that appeared in the dormant state. Large and abundant lipid bodies were present in the active cambium cells. No differences were observed in the nucleus and mitochondria between active and dormant states. Conclusions: The number of cell layers and the morphology of cells in the cambium enabled us to define whether it is dormant or active. In active and dormant states, some dissimilarities in the shape and intensity of organelles were observed during the growth-to-dormancy transition, emphasizing the importance of stress protection in the tree's adaptive response to overwintering. Endoplasmic reticulum, starch granules, vacuoles, and lipid droplets showed great distinguishing characteristics between active and dormant cambium cells in white spruce. [ABSTRACT FROM AUTHOR]

Losses in forest productivity through poor soil management are typically evaluated by changes in crop production, but conflicting growth responses among co-occurring species can challenge criteria for sustainability. In this study, we evaluate species response to compaction and organic matter removal by contrasting the growth and foliar attributes at age 20 of lodgepole pine ( Pinus contorta Dougl. ex Loud.) and hybrid white spruce ( Picea glauca × engelmannii (Moench) Voss) in sub-boreal forests of central British Columbia. Bole volume increment was stable for lodgepole pine, averaging a 6 % difference across treatments. White spruce, in contrast, often had large growth increases on forest floor retained - compacted plots (average of 63 % gain) but reductions in productivity on the forest floor removed - compacted plots (40 % decline). Foliar nitrogen concentrations converged across treatments for both species, despite the removal of over 50 % of the site N capital. Some key differences in species traits were suggested by foliar δ15N, attributed to deeper rooting of pine, and foliar δ13C, indicating possibly higher CO2 assimilation potential for spruce. The implication is that standard metrics of sustainable forestry are contingent upon tree autecology and that comprehensive assessments of land management require a measure of collective species response. [ABSTRACT FROM AUTHOR]