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Future warming may alter plant stress at high‐elevation treelines and forests, thereby changing plant–plant interactions. The relative importance of competition and facilitation may depend on the degree of resource or physical stress. According to the stress gradient hypothesis (SGH), physical stress on trees is more important at cold high elevations where facilitation predominates, and less important at low elevations where competition is the main interaction. Our goals were to investigate whether plant–plant interactions along elevational gradients corresponded to those predicted by the SGH, and to assess the effects of increasing temperatures on the growth of conifer seedlings in tundra–forest ecosystems in interior Alaska, USA. We established sites along two elevational gradients: one in tundra (four sites, 550–1170 m) and one in forest (three sites, 210–760 m). A field warming and neighbor removal experiment was conducted using transplanted seedlings of white spruce (Picea glauca). After three growing seasons, regardless of elevation, spruce seedling biomass was approximately 20% lower in seedlings with neighbor plants compared to those without neighbors. Therefore, there was no evidence for greater facilitation at high elevations across either tundra or forest elevational gradient. Seedlings in forest sites increased their shoot‐to‐root ratios and relative growth rates in height, suggesting competition for light, but this was not seen for seedlings growing in tundra sites. When warmed by greenhouses, seedling growth was stimulated in forests, especially at high elevations. In contrast, at tundra sites, warmed seedlings grew less than controls and the seedlings at high elevations showed water stress. Our results suggest that the resource stresses of low light and water availability are more important drivers of plant–plant interactions than the physical stress imposed by low temperature, possibly due to warming in the past 50 yr in this region. Further warming may increase growth of seedlings in forests, but is likely to decrease it in open tundra habitats, thereby slowing the upward movement of treeline in the near future. [ABSTRACT FROM AUTHOR]

With climate change, favorable growing conditions for tree species are shifting northwards and to higher altitudes. Therefore, local populations are becoming less adapted to their environment. Assisted migration is one of the proposed adaptive measures to reduce the vulnerability of natural populations and maintain forest productivity. It consists of moving genetic material to a territory where future climate conditions correspond to those of its current location. Eight white spruce (Picea glauca [Moench] Voss) seed sources representing as many seed orchards were planted in 2013 at three forest sites simulating a south-north climatic gradient of 1.7°C in Québec, Canada. The objectives were to (1) evaluate the morpho-physiological responses of the different seed sources and (2) determine the role of genetic adaptation and physiological plasticity on the observed variation in morpho-physiological traits. Various seedling characteristics were measured, notably height growth from nursery to the fourth year on plantation. Other traits such as biomass and carbon allocation, nutritional status, and various photosynthetic traits before bud break, were evaluated during the fourth growing season. No interaction between sites and seed sources was observed for any traits, suggesting similar plasticity between seed sources. There was no change in the rank of seed sources and sites between years for height growth. Moreover, a significant positive correlation was observed between the height from the nursery and that after 4 years in the plantation. Southern seed sources showed the best height growth, while optimum growth was observed at the central site. Juvenile height growth seems to be a good indicator of the juvenile carbon sequestration and could serve as a selection criterion for the best genetics sources for carbon sequestration. Vector analysis showed no nitrogen deficiency 4 years after planting. Neither seed sources nor planting sites had a significant effect on photosynthesis before bud break. The observed results during the establishment phase under different site conditions indicate that southern seed sources may already benefit from assisted migration to cooler climatic conditions further north. While northern seed sources are likely to benefit from anticipated local global warming, they would not match the growth performance of seedlings from southern sources. [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]

Climate change is expected to increase winter temperatures in boreal climates. White spruce ( Picea glauca (Moench) Voss) is vulnerable to spring frost damage due to its habit of early budbreak, which may be exacerbated or lessened with increasingly warm winters at its southern range edge. We tested the effects of episodic warming during the quiescent stage on budbreak time and growth of seven seed sources grown in a common garden setting in Minnesota, USA. Treatment plots were warmed with infrared lamps for 4 days each in February, March, or February and March to simulate a midwinter thaw. Control plots for each treatment and an overall control were included for comparison. Trees warmed in February experienced a slight delay in spring budbreak, but differences in budbreak time were generally not significant. Terminal growth was significantly and negatively correlated with time of budbreak but not with time to growth cessation. Our results suggest that white spruce is relatively resilient to the effects of intermittent warming but that warming early in the season may delay budbreak time, which is expected to reduce terminal growth. [ABSTRACT FROM AUTHOR]

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high-latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [ Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down-regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 (Nmass), 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 (Amax), stomatal conductance (gs) and mesophyll conductance (gm). Total dry mass, but not height growth, was strongly correlated with latitude of seed origin (SO) and associated climate variables. Amax, gs, gm 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 (Vcmax), maximum rates of electron transport (Jmax), SLA, Nmass, 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. [ABSTRACT FROM AUTHOR]

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 CO2over two growing seasons. There was positive effect of warmer temperatures and higher CO2on some growth variables but no significant family × treatment interactions. Instantaneous water use efficiency was the only physiological trait that was affected positively by the CO2treatment, showing a 51% increase that was consistent across families. [ABSTRACT FROM PUBLISHER]

CO2–light interactions can influence the competition among boreal plants, but are poorly understood. We investigated the effect of such interactions on the growth and biomass of 1-year-old black spruce (Sb) (Picea mariana (Mill.) BSP) and white spruce (Sw) (Picea glauca (Moench) Voss) grown with CO2 concentrations ([CO2]) of 360 and 720 µmol·mol–1 under 30%, 50%, and 100% light, in greenhouses. There were significant two-way and three-way interactions. Root collar diameter (RCD) of Sw decreased with decreasing light, while in Sb, there was no significant difference in RCD for plants grown under 50% or 30% light. Height was greater for plants grown under 100% light than if the plants were shaded. Elevated [CO2] increased RCD by 33% and enhanced stem volume by 67%, 98%, and 84% under 100%, 50%, and 30% light, respectively. The CO2 enhancement of total biomass was relatively higher under lower light, and greater for Sb than Sw. Elevated [CO2] decreased specific leaf area under 50% light only. Root mass was generally higher under 100% light than when shaded. Elevated [CO2] increased the root mass of Sb under 100% light, but decreased it under 30% light. Elevated [CO2] decreased the shoot/root ratio under 100% light, but increased it under 30% light. Our data suggest that raising [CO2] will likely increase species competitiveness under low light conditions, and that the increase will be greater in species that are relatively shade tolerant. [ABSTRACT FROM AUTHOR]

Aim The objectives of the study are: (1) to evaluate the dynamics of the maritime tree line and forest limit of white spruce, Picea glauca, within the dual framework of primary succession induced by the rapid post-glacial land emergence on the eastern coast of Hudson Bay and the impacts of recent and past climate changes; and (2) to determine the time lapse between land emergence and seedling, tree, and forest establishment in the context of the primary chronosequence occurring on rising, well-drained sandy beaches and terraces. Location The study area was located on the eastern coast of Hudson Bay (56°20′ N, 76°32′ W) in northern Québec, Canada. Methods We evaluated the colonization dynamics of white spruce as seedlings, tree-line trees and primary-forest trees at eight sites distributed along a 200-km latitudinal gradient based on a mean land emergence rate of 1.2 m century−1. A 30-m wide by 140–300-m long quadrat was positioned at random at the centre of each site. The elevation above sea level, position and age of all individuals of spruce present in the quadrat areas were determined, and the soils of each chronosequence were described. Results The main stages of primary succession along the emerging coast were common to all the sites, regardless of latitude, but occurred at different elevations above sea level (a.s.l.). White spruce seedlings colonized near-shore beaches 2 m a.s.l., whereas the tree line and forest limit tended to form only at about 3–4 m and 4–8 m a.s.l., corresponding approximately to 180–825 years and 310–1615 years after land emersion, respectively. White spruce establishment at the tree line occurred about 50 years ago. Climatic conditions at this time were probably more favourable to tree colonization than when the species established at the forest limit. Soil formation was influenced primarily by distance from the seashore and elevation above sea level, with podzolization being accelerated by white spruce cover. Main conclusions The current tree-line and forest-limit positions on the rising coast of eastern Hudson Bay correspond to ecological limits established during the course of primary succession within a context of changing climatic conditions. The recent establishment of trees at the tree line and forest limit at relatively old coastal sites is associated with warmer conditions over the last 100 years. Although white spruce was present nearby, coastal sites were devoid of trees before the 20th century. [ABSTRACT FROM AUTHOR]

From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [ Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open-top chambers (OTCs) distributed equally between opposing north and south-facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1. 8°C and annual growing degree-days by one-third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1. 0°C warmer and the snow-free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem-II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure-related differences. Our results suggest that the growth of seedlings on north-facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south-facing slopes appears limited by winter desiccation and cold-induced photoinhibition. [ABSTRACT FROM AUTHOR]