Irregular Heartwood Formation in Maritime Pine (Pinus pinaster Ait):Consequences for Biomechanical and Hydraulic Tree Functioning

As the proportion of sapwood (SW) transformed into heartwood(HW) is irregular both radially and longitudinally in trunksof Maritime pine (Pinus pinaster Ait.), it has been suggestedthat HW formation is a developmental process, regulated internallywithin the tree. In trees where stem growth is eccentric dueto stem lean or wind action, the number of annual growth ringsof SW transformed into HW is greater on the compressed sideof the tree. To determine the contribution to bending stiffness,if any, of this prematurely formed HW, four point bending testswere carried out on fresh HW and SW samples taken from the samegrowth ring, or neighbouring growth rings, at different cross-sectionalpositions at a height of 2 m from six 52-year-old Maritime pines.The mean (±s.e.) modulus of elasticity (E) was 7.6 ±0.3 GPa (longitudinal direction) for all samples. No significantdifferences in E were found between HW and SW; thus HW doesnot play a significant mechanical role in bending stiffness.To test a second hypothesis that early HW formation on the compressedside of trees may maintain a constant, optimal volume of SWaround the tree, the Pipe Model Theory was applied to 12 52-year-oldleaning Maritime pines (angle of lean varied from 0–22°).The surface area (S) of the SW was determined at different heightsup the trunk and correlated with crown surface area (S_crown).Regressions between S_SWandS_crown were highly significant, therebysupporting the theory that HW formation and extension is controlledinternally in Maritime pine. HW formation in Maritime pine thenserves to maintain an optimal proportion of functional SW whichis an important criterion for survival in a species often subjectedto severe drought for long periods. Copyright 2001 Annals ofBotany Company Heartwood, sapwood, bending test, Pipe Model Theory, Pinus pinaster     

Above- and Below-ground Production of Young Scots Pine (Pinus sylvestris L.) Trees after Three Years of Growth in the Field under Elevated CO2

Scots pine (Pinus sylvestris L.) seedlings were grown for 3years in the ground in open top chambers and exposed to twoconcentrations of atmospheric CO₂(ambient or ambient + 400 µmol mol^-1) without addition of nutrients and water. Biomassproduction (above-ground and below-ground) and allocation, aswell as canopy structure and tissue nitrogen concentrationsand contents, were examined by destructive harvest after 3 years.Elevated CO₂increased total biomass production by 55%, reducedneedle area and needle mass as indicated, respectively, by lowerleaf area ratio and leaf mass ratio. A relatively smaller totalneedle area was produced in relation to fine roots under elevatedCO₂. The proportion of dry matter in roots was increased byelevated CO₂, as indicated by increased root-to-shoot ratioand root mass ratio. Within the root system, there was a significantshift in the allocation towards fine roots. Root litter constituteda much higher fraction of fine roots in trees grown in the elevatedCO₂than in those grown in ambient CO₂. Growth at elevated CO₂causeda significant decline in nitrogen concentration only in theneedles, while nitrogen content significantly increased in branchesand fine roots (with diameter less than 1 mm). There were nochanges in crown structure (branch number and needle area distribution).Based upon measurements of growth made throughout the 3 years,the greatest increase in biomass under elevated CO₂took placemainly at the beginning of the experiment, when trees grownin elevated CO₂had higher relative growth rates than those grownunder ambient CO₂; these differences disappeared with time.Symptoms of acclimation of trees to growth in the elevated CO₂treatmentwere observed and are discussed. Copyright 2000 Annals of BotanyCompany Elevated CO₂, Pinus sylvestris, biomass production, allocation, fine roots, root litter, crown structure, nitrogen, C/N ratio     

Modelling and Measuring Transpiration from Scots Pine with Increased Temperature and Carbon Dioxide Enrichment

Starting in 1996, individual trees of Scots pine (Pinus sylvestrisL.) aged 30 years, were grown in closed-top chambers and exposedto either normal ambient conditions (CON), elevated CO₂(approx.700 µmol mol^-1; Elev. C), elevated temperature (approx.2 °C and approx. 6 °C above the outside ambient temperatureduring the ‘growing season’ and ‘off season’,respectively; Elev. T) or a combination of elevated CO₂and warmertemperature (Elev. CT). Sap flow was monitored simultaneouslyby the constant-power heat balance method in a total of 16 trees,four for each treatment, over a 32 d period in summer 1998 (afterthe completion of needle expansion and branch elongation). Toquantify the contributions of crown and physical environmentalvariables to total crown transpiration, a ‘sun/shade model’was developed and used to partition the changes in transpirationto different sources. The results of the sap flow measurementsindicate that (1) total daily sap flow (E_tree.d) varied from0.15–3.41 kg per tree; (2) the treatment effect on E_tree.ddependedgreatly on the weather conditions; (3) the cumulative E_tree.dforthe 32 d dropped significantly by 22% relative to CON (P =0.038)under Elev. C and increased significantly by 21% (P =0.043)and 16% (P =0.048) under Elev. T and Elev. CT, respectively.In general, the modelled transpiration gave good agreement withthe sap flow results. The model computations showed that, ona typical sunny day in summer, the effect of treatment on crownstomatal conductance was responsible for approx. 80% of thechange inE_tree.d , while the increase in needle area and theeffect on total radiation absorption contributed only a smallpercentage. Furthermore, sunlit needles were responsible forover 60% of change in transpiration. The effect of the treatmentson E_tree.dwas larger at high temperature and vapour pressuredeficit but was not sensitive to incident daily radiation. Copyright2000 Annals of Botany Company Transpiration model, sap flow, CO₂and temperature elevation, environment-controlled chamber, Pinus sylvestris L.     

Understanding 3D structural complexity of individual Scots pine trees with different management history

Tree functional traits together with processes such as forest regeneration, growth, and mortality affect forest and tree structure. Forest management inherently impacts these processes. Moreover, forest structure, biodiversity, resilience, and carbon uptake can be sustained and enhanced with forest management activities. To assess structural complexity of individual trees, comprehensive and quantitative measures are needed, and they are often lacking for current forest management practices. Here, we utilized 3D information from individual Scots pine (Pinus sylvestris L.) trees obtained with terrestrial laser scanning to, first, assess effects of forest management on structural complexity of individual trees and, second, understand relationship between several tree attributes and structural complexity. We studied structural complexity of individual trees represented by a single scale‐independent metric called “box dimension.” This study aimed at identifying drivers affecting structural complexity of individual Scots pine trees in boreal forest conditions. The results showed that thinning increased structural complexity of individual Scots pine trees. Furthermore, we found a relationship between structural complexity and stem and crown size and shape as well as tree growth. Thus, it can be concluded that forest management affected structural complexity of individual Scots pine trees in managed boreal forests, and stem, crown, and growth attributes were identified as drivers of it.     

Modelling the distribution of diameter growth along the stem in Scots pine

This paper presents an empirical model for the distribution of diameter growth along the stem in Scots pine (Pinus sylvestris L.) and for the consequent stem form over time. First, the distribution of annual mass growth in the stem is determined as a function of the total annual growth in stem mass, current stem mass and the distribution of the latter along the stem. Second, the distribution of diameter growth is obtained by converting the fraction of annual growth in the stem mass at a given height in the stem into the thickness of the annual ring at the same height. Application of the model to Scots pine data sets including both young and mature trees not used in parameter estimation showed that the model was capable of reconstructing the distribution of diameter growth from the stem butt to the apex and from the pith to the stem surface at any height in the stem in both young and mature trees. The resulting empirical model was also linked to a physiological, process-based model in order to study its performance in a simulated stand. Simulations representing trees grown in unthinned and thinned Scots pine stands with trees of different status (from dominant to suppressed) showed that the response in tree growth to thinning in terms of the distribution of diameter growth along the stem was quite realistic relative to measured data.     

Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Bursts of CO2 released during freezing offer a new perspective on avoidance of winter embolism in trees

MethodsCO₂ efflux measurements were conducted during freezing experiments for saplings of three Scots pine (Pinus sylvestris) and three Norway spruce (Picea abies) trees under laboratory conditions, and the magnitudes of the freezing-related bursts of CO₂ released from the stems were analysed using a previously published mechanistic model of CO₂ production, storage, diffusion and efflux from a tree stem. The freezing-related bursts of CO₂ released from a mature Scots pine tree growing in field conditions were also measured and analysed.ConclusionsAll gases dissolved in the xylem sap are not trapped within the ice in the stem during freezing, as has previously been assumed, thus adding a new dimension to the understanding of winter embolism formation. The conduit water volume not only determines the volume of bubbles formed during freezing, but also the efficiency of gas efflux out of the conduit during the freezing process.     

Growth and Nutrient Retranslocation in Needles of Radiata Pine in Relation to Nitrogen Supply

The effects of N application on tree growth and the retranslocationof N, P, and K from young needles to new growth were examinedin young radiata pine (Pinus radiata D. Don) trees. Nitrogen fertilization increased the number and size of needles,rates of shoot production, stem volume growth and tree biomass.Foliar N and P contents (µg per needle) fluctuated ina cyclic fashion with prominent phases of accumulation, retranslocationand replenishment. The patterns of these fluctuations in controland N-fertilized trees were similar, although the fluxes ofN, P and K in and out of needles were increased by N fertilization.Greater translocation (g per tree) of N and K from needles ofN fertilized trees occurred because fertilization increasedthe needle weight and the proportion of N and K retranslocatedfrom individual needles. Nitrogen fertilization increased theretranslocation of P largely as a result of higher needle mass.Trees supplied with more than adequate amounts of P in the soilretranslocated up to 58 per cent of the initial pool of P fromyoung needles. The periods of high retranslocation coincidedwith periods of high concentrations of soil mineral N and withshoot production. Conversely, the periods of rapid replenishmentof N and P into the needles coincided with the time of slowshoot growth and low concentration of soil mineral N. The growthrate of trees, rather than the availability of nutrients inthe soil was the main factor controlling retranslocation. For radiata pine, retranslocation from needles is not a mechanismspecific for coping with low soil fertility. It seems to bea mechanism which enhances the nutrient supply to apical growingpoints, especially during periods of flushing. Pinus radiata, nitrogen supply, shoot growth, nutrient fluctuations and retranslocation, nutrient use and adaptation     

Boron Mobility in Two Coniferous Species

In contrast to earlier beliefs, it is now known that boron (B)can be retranslocated complexed with sugar alcohols in someplant species. Conifers had been thought not to translocatesugar alcohols in the phloem. However, 1 d after applying¹⁰Benriched boric acid to shoots of Scots pine and Norway spruceseedlings, we found increases in both the amount and proportionof¹⁰B in the root systems in both species. We conclude thatB is translocated in the phloem from shoots to roots in spruceand pine, and therefore it is possible that these species retranslocateB. Copyright 2000 Annals of Botany Company Scots pine, Pinus sylvestris, Norway spruce, Picea abies, conifers, boron retranslocation, roots, stable isotopes, sugar alcohols, boron complexes, mineral nutrition, forest trees     

Shoot Elongation in Scots Pine: Diurnal Variations and Response to Temperature

Kanninen, M. 1985. Shoot elongation in Scots pine: Diurnal variationsand response to temperature.—J. exp. Bot. 36: 1760–1770. Time series analysis is used to study the diurnal variationsin the rate of shoot elongation of Scots pine (Pinus sylvestrisL.) seedlings, and its response to temperature. The shoot elongation of three 5-year-old Scots pine seedlingswas measured at 2 h intervals over a 12 d period in June 1979.Ambient temperature, measured with a thermocouple, was continuouslyrecorded during the same period. The measurements were carriedout in the field. Time series analysis of the data was carried out by estimatedauto-correlation and partial auto-correlation functions. A multiplicative‘seasonal’ model was applied to both the input andoutput series to ‘prewhiten’ the data set. Serialcross-correlation analysis of the prewhitened series was usedto identify the structure of the transfer function model betweentemperature and growth rate. There was a phase shift between the air temperature and shootextension. The cross-correlation function peaked at 2 h lagvalue. The influence of temperature on the growth rate was adequatelydescribed by a first-order transfer function model. The characteristicfeature of the model was a time constant of 3.3 h and a basetemperature of 6.3 °C for the response of shoot elongationto temperature. Key words: Scots pine, Pinus sylvestris L., shoot elongation, temperature response, time-series analysis     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Atmospheric ozone affects carbohydrate allocation and winter hardiness of Pinus halepensis (Mill.)

Fumigation of 2 year-old Aleppo pine (Pinus halepensis Mill.)trees with episodes of O₃ (up to 120 nl l^– as comparedto 25 nl l^–) throughout two summer seasons produces tallerplants with greater stem diameters but reduced root:shoot ratios.Both light and electron microscopy of current year needles carriedout at mid-summer reveal extensive accumulations of starch,particularly in the endodermis, and crushing of the phloem sievecells. Parallel measurements of starch levels also reveal anon-mobilizable component. All these features probably indicatethat episodes of high summer O₃ detrimentally affect the normalability of Aleppo pines to withstand severe water stress andthis may account for their reduced vitality in Mediterraneanregions. By autumn, however, the amounts of starch are similarto those in low O₃-grown trees and no evidence of the phloemsieve tube crushing remain although plastids do show increasednum bers of plastoglobull. Similar summer high level O₃-fumigatedneedles also show enhanced winter hardiness in the autumn whichis unusual for conifers. Key words: Pinus halepensis, Aleppo pine, amyloplasts, ozone, plastoglobuli, starch     

An Examination of the Differences in Dry Matter Production shown by Some Progenies of Pinus sylvestris L.

A study was carried out to determine the extent of the variationin growth rate between several selected progenies of Scots pine{Pinus sylvestris L.) and the physiological characteristicsunderlying such variation, using growth analysis and directmeasurement of photosynthetic rates. Differences in relativegrowth rate (RGR) could be demonstrated for first and secondyear seedling progenies, and these largely reflected differencesin net assimilation rate (NAR). The relative performance ofthe progenies in different experiments was not always the same,although the highest RGR was shown by the same progeny in bothfirst-year and second-year seedlings. For most progenies theobservations were extended to 14-year-old trees growing in fieldtrials established by the Forestry Commission. Comparisons ofthe relative height growth rates (RHGR) of these trees showedno significant differences between the progenies, apparentlybecause the growth rates were limited by the site conditions.Thus, the size differences at the end of the fourteenth yearprobably still reflected differences in RGR which occurred duringthe seedling stages in the nursery. The significance of theseobservations for the assessment of performance in tree breedingprogrammes is discussed.     

Isolation and Purification of Mitochondrial Mn-Superoxide Dismutase from the Gymnosperm Pinus sylvestris L.

Manganese superoxide dismutase (Mn-SOD; EC 1.15.1.1 [EC] ) was purifiedfrom germinating seeds of Scots pine (Pinus sylvestris L.) 3days after the start of imbibition. The purification scheduleincluded (NH₄)₂SO₄ fractionation, anion-exchange and hydrophobic-interactionchromatographies and chromatofocusing. Purified Mn-SOD had anapparent specific activity of 4,130 McCord-Fridovich units (mgprotein)^–1. The molecular mass of the holoenzyme was estimatedto be 91 kDa by size-exclusion chromatography, and a molecularmass of 23 kDa was determined by SDS-PAGE. However, isoelectricfocusing demonstrated that the purified enzyme consisted ofthree similarly migrating isoforms, with isoelectric pointsof approximately 6.5. NH₂-terminal amino acid sequencing ofpurified Mn-SOD revealed no differences among the three isoforms.The comparison of the first 32 NH₂-terminal amino acids withsequences of NH₂-terminal amino acids of Mn-SODs from angiospermsreflected the phylogenetic distances between Scots pine, whichis a gymnosperm, and angiospermic species. Cell fractionationsuggested the mitochondrial localization of Mn-SODs and no evidencefor glyoxysomal localization was found. Mn-SOD activity wasabsent from dry seeds. It was detectable at a considerable levelafter imbibition for 24 h, and it was again absent from 3-week-oldseedlings. (Received February 8, 1994; Accepted May 24, 1994)     

Variability of some phenolic acids in phloem of 1-year-old shoots of Scots pine trees growing with Heterobasidion annosum (Fr.) Bref.

 HPLC chromatographic analyses of some phenolic acids in phloem of 1-year-old shoots sampled from 32 trees of eight Polish provenances of Scots pine (Pinus sylvestris L.) growing under conditions of annosum root [Heterobasidion annosum (Fr.) Bref.] are discussed. Considerable quantitative and qualitative differentiation was found among individual trees. The variability of trees was estimated with regard to the level of phenolic acids and correlations were established in order to assess the character of their joint occurrence in shoot phloem. In view of pathogen presence, the content of phenolic acids varies between individuals depending upon the genotype of pine, the stage of development of the disease and upon the effect of tree growth conditions. Received: 23 April 1997 / Accepted: 12 September 1997     

Seasonal changes of pinosylvin distribution in the sapwood/heartwood boundary of Pinus sylvestris

Changes in pinosylvin concentration and distribution across the sapwood/heartwood boundary were studied on Scots pine (Pinus sylvestris L.) tree stems to detect seasonal activity in heartwood formation. Pinosylvin concentrations were measured with FT-(NIR) Raman spectroscopy for a total of 96 trees equally distributed on 16 different sampling occasions. In another experiment, cores were sampled every month from six individual Scots pine trees from June to October and analysed for pinosylvin. There was a great between-tree variation in concentration and spatial distribution of pinosylvin. No seasonal trend in the distribution pattern or concentration of pinosylvin was found. Therefore, the results indicate that there is no specific period during the year when heartwood is formed. Received: 12 February 1998 / Accepted: 3 May 1999     

Connections between climatic variables and the growth and needle dynamics of Scots pine (Pinus sylvestris L.) in Estonia and Lapland

The relationships between climatic variables and Scots pine (Pinus sylvestris L.) growth and needle dynamics were studied in three stands in Estonia and in four stands located near the northern timberline in Lapland. The trees sampled in Estonia had low correlations with the analysed climatic variables (air temperature, precipitation and indices of atmospheric circulation). Moreover, the weak cross-correlation of the time-series of the Estonian sample trees indicated that Scots pine is affected mainly by local factors in that region. In Lapland, however, height increment and needle production correlated strongly among trees within a stand (mean r=0.45 and 0.46, respectively) and between stands (r=0.32 and 0.37). Radial increment also showed a high inter-correlation among the trees within a stand in Lapland (r=0.45). Both height increment and needle production were strongly influenced by the temperature regime of the previous summer in Lapland (mean r=0.64 and 0.64, respectively). Radial increment was correlated with the mean July temperature of the current year (mean r=0.29). The correlations between the indices of atmospheric circulation and tree attributes were weak, while the strongest correlation was between the Ponta Delgada NAO index (PD–NAO) and height increment and needle production in Lapland. Height increment, needle production and radial increment have increased since the 1990s in the trees growing in Lapland. This may indicate a positive effect of climate warming on tree growth in Lapland. In Estonia, where climatic conditions do not limit tree growth, the climate warming seems not to directly influence the growth and needle dynamics of Scots pine.     

Towards a new approach for dendroprovenancing pines in the Mediterranean Iberian Peninsula

Dendroprovenancing studies frequently use site chronologies to identify the origin of archaeological and historical timber. However, radial growth (tree-ring width, TRW) of tree species is influenced by both local and regional climate scales. Here we investigate how the use of annually-resolved Blue Intensity (BI) measurements can enhance dendroprovenancing precision of black pine (Pinus nigra Arn.) and Scots pine (P. sylvestris L.) on the Iberian Peninsula. Principal Component Gradient Analyses (PCGA) was used to assess geographical patterns of annual variation in different TRW and BI proxies of pine trees from two mountain ranges in the Central System and Andalusia in Spain. Local climate-growth relationships were quantified to identify underlying causes of identified groups with diverse growth patterns. Two distinct elevational groups were observed when performing PCGA on latewood BI time series with the response to summer drought as the main factor causing the differences. Both P. nigra and P. sylvestris BI time series were found to be more related to summer drought at low-elevation sites showing an increase in sensitivity at lower latitudes. PCGA of TRW time series allowed to discriminate between trees from Andalusia and Central System within the elevation groups. February and October temperatures were found to be the main climatic factors causing the differences in TRW time series among the high- elevation sites, whereas for low-elevation trees it was the average winter temperature influencing TRW. A subsequent leave-one-out analyses confirmed that including latewood BI time series improves the precision of dendroprovenancing of pine wood in the Iberian Peninsula.     

Effect of water availability and fertilization on water status,growth, vigour and the resistance of Scots pine to fungal mass inoculation with Ophiostoma ips

Abstract

The effect of water and nutrient availability on the performance of Scots pine (Pinus sylvestris L.) against Ophiostoma ips (Rumb.), a bark beetle-associated phytopathogenic blue-stain fungus, was investigated. Field-grown trees were subjected for 18 months to water-stress and/or fertilization, and the effects of such treatments on the needle nutrient status, tree vegetative growth and vigour were examined. At the end of the experimental period, the trees were mass-inoculated (800 inocula m^?2) with the fungus, and the relationship between resource availability and tree performance against pathogen attack was also tested. Predawn shoot water potential (Ψ_PD) of irrigated trees was significantly higher than that of water-stressed trees, and fertilized trees had a significantly lower C/N ratio. The Ψ_PD values and needle nitrogen content suggest that resource-limited trees were under moderate stress. Improved nutrient availability significantly increased tree growth and tree vigour. However, no evidence for an effect of improved nutrient availability on tree fungal resistance was found in our study.     

Temperature and soil fertility as regulators of tree line Scots pine growth and survival—implications for the acclimation capacity of northern populations

The acclimation capacity of leading edge tree populations is crucially important in a warming climate. Theoretical considerations suggest that adaptation through genetic change is needed, but this may be a slow process. Both positive and catastrophic outcomes have been predicted, while empirical studies have lagged behind theory development. Here we present results of a 30‐year study of 55,000 Scots pine (Pinus sylvestris) trees, planted in 15 common gardens in three consecutive years near and beyond the present Scots pine tree line. Our results show that, contrary to earlier predictions, even long‐distance transfers to the North can be successful when soil fertility is high. This suggests that present northern populations have a very high acclimation capacity. We also found that while temperature largely controls Scots pine growth, soil nutrient availability plays an important role—in concert with interpopulation genetic variation—in Scots pine survival and fitness in tree line conditions. These results suggest that rapid range expansions and substantial growth enhancements of Scots pine are possible in fertile sites as seed production and soil nutrient mineralization are both known to increase under a warming climate. Finally, as the ontogenetic pattern of tree mortality was highly site specific and unpredictable, our results emphasize the need for long‐term field trials when searching for the factors that control fitness of trees in the variable edaphic and climatic conditions of the far North.