Responses in growth and emissions of biogenic volatile organic compounds in Scots pine, Norway spruce and silver birch seedlings to different warming treatments in a controlled field experiment.

We investigated the responses in growth and emissions of biogenic volatile organic compounds (BVOCs) in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies [L.] Karst.) and silver birch (Betula pendula Roth) seedlings to a simulated climate warming of +2 °C (T2) and +4 °C (T4), compared to the ambient conditions, during two growing seasons (2019-2020) in a controlled field experiment in central Finland. In all seedlings, height was measured weekly. Diameter was measured continuously for one seedling from each tree species per plot. For shoot and root biomass measurements, half of the seedlings were harvested at end of the first growing season and the rest at the end of the second growing season. Foliage BVOC emission rates were measured at the end of the second growing season. Biomass, height, and diameter growth of silver birch did benefit the most from warming in both growing seasons. In the Scots pine and Norway spruce seedlings, height and diameter growth increased with increasing temperature in the second growing season, more so in Scots pine. Overall, the shoot and root biomass of conifer seedlings increased with increasing temperature. In the conifer seedlings, warming increased biomass and diameter growth more than height growth, due to their predetermined height growth pattern. The warming increased BVOC emissions more clearly in silver birch, whilst the BVOC emissions were in conifers less sensitive to temperature variation. Based on our findings, silver birch seedlings could be expected to benefit the most from warmer growing conditions and Norway spruce the least.

Accumulation of phenolics and growth of dioecious Populus tremula (L.) seedlings over three growing seasons under elevated temperature and UVB radiation.

Accumulation of secondary metabolites may exhibit developmentally regulated variation in different plant organs. Moreover, prevailing environmental conditions may interact with development-related variations in plant traits. In this study, we examined developmentally regulated variation in phenolic accumulation in the twigs of dioecious Populus tremula (L.) and how the effects of elevated temperature and ultraviolet B (UVB) radiation on growth and phenolics accumulation varied as the plants get older. In an open-field experiment, six female and six male genotypes were exposed to single and combined elevated temperature and UVB radiation treatments for three consecutive growing seasons. The concentrations of low molecular weight phenolics and condensed tannins did not show age-dependent variation in the twigs. In temperature-treated plants, diameter growth rate decreased, and concentration of condensed tannins increased as plants aged; there were no cumulative effects of elevated UVB radiation on growth and phenolic accumulation. Females maintained a higher concentration of low molecular weight phenolics throughout the experimental period; however, growth and phenolic concentration did not vary over time in females and males. Our results suggest that phenolic accumulation in perennial plants may not necessarily always exhibit age-dependent variation and the effects of elevated temperature on growth and phenolic may diminish as plants get older.

Climate change induces multiple risks to boreal forests and forestry in Finland: A literature review.

Climate change induces multiple abiotic and biotic risks to forests and forestry. Risks in different spatial and temporal scales must be considered to ensure preconditions for sustainable multifunctional management of forests for different ecosystem services. For this purpose, the present review article summarizes the most recent findings on major abiotic and biotic risks to boreal forests in Finland under the current and changing climate, with the focus on windstorms, heavy snow loading, drought and forest fires and major insect pests and pathogens of trees. In general, the forest growth is projected to increase mainly in northern Finland. In the south, the growing conditions may become suboptimal, particularly for Norway spruce. Although the wind climate does not change remarkably, wind damage risk will increase especially in the south, because of the shortening of the soil frost period. The risk of snow damage is anticipated to increase in the north and decrease in the south. Increasing drought in summer will boost the risk of large-scale forest fires. Also, the warmer climate increases the risk of bark beetle outbreaks and the wood decay by Heterobasidion root rot in coniferous forests. The probability of detrimental cascading events, such as those caused by a large-scale wind damage followed by a widespread bark beetle outbreak, will increase remarkably in the future. Therefore, the simultaneous consideration of the biotic and abiotic risks is essential.

Responses in growth and phenolics accumulation to lateral bud removal in male and female saplings of Populus tremula (L.) under simulated climate change.

The responses in growth and defense after tissue damage are highly variable in plants depending on species, damaged-tissue type and the intensity of damage. The prevailing abiotic conditions can also influence these responses. In this study, our aim was to examine how the removal of lateral vegetative buds affects the growth and accumulation of phenolics in saplings of the dioecious Populus tremula grown under simulated climate change. For three growing seasons, the saplings were grown under ambient conditions (control), elevated temperature (+2°C) and elevated UV radiation (30%) (UVB and UVA as its control), or a combination of these. In the fourth growing season, all saplings were grown under ambient conditions. The bud removal was performed twice - in summer and autumn - in the third year. The responses of growth and the accumulation of phenolics to the bud removal were measured at the end of the fourth growing season. Removal of 5% of the lateral buds resulted in higher leaf, stem and total plant biomass in both sexes of P. tremula saplings, compared to intact plants. The effects were greater in the temperature-treated plants, especially in the temperature-treated females. The concentrations of flavonoids and condensed tannins were higher in the bud-removed individuals. The concentration of condensed tannins was also higher in the males than in the females, opposite to the concentration of phenolic acids. There was no significant interaction between bud removal and UVB treatment on either growth or phenolics. Our results suggest that plants can allocate resources to both growth and defense simultaneously in response to tissue loss, and that global warming can modify the responses to some extent.

Effect of increased wood harvesting and utilization on required greenhouse gas displacement factors of wood-based products and fuels.

A displacement factor (DF) may be used to describe the efficiency of using wood-based products or fuels instead of fossil-based ones to reduce net greenhouse gas (GHG) emissions. However, the DFs of individual products and their production volumes could not be used alone to evaluate the climate impacts of forest utilization. For this reason, in this study we have developed a methodology to assess a required displacement factor (RDF) for all wood products and bioenergy manufactured and harvested in a certain country in order to achieve zero CO2 equivalent emissions from increased forest utilization over time in comparison with a selected baseline harvesting scenario. Input data for calculations were produced with the simulation model, Monsu, capable of predicting the carbon stocks of forests and wood-based products. We tested the calculations in Finnish conditions in a 100-year time horizon and estimated the current average DF of manufactured wood-based products and fuels in Finland for the interpretation of RDF results. The results showed that if domestic wood harvesting will be increased by 17-33% compared to the basic scenario, the RDF will be 2.0 to 2.4 tC tC-1 for increased wood use in 2017-2116. However, the estimated average DF of manufactured wood-based products and fuels currently in Finland was less than 1.1 tC tC-1. The results indicate strongly that the increased harvesting intensity from the current situation would represent a challenge for the Finnish forest-based bioeconomy from the viewpoint of climate change mitigation. For this reason, there is an immediate need to improve reliability and applicability of the RDF approach by repeating corresponding calculations in different circumstances and by improving estimations of DFs on country levels.

Are forest disturbances amplifying or canceling out climate change-induced productivity changes in European forests?

Recent studies projecting future climate change impacts on forests mainly consider either the effects of climate change on productivity or on disturbances. However, productivity and disturbances are intrinsically linked because 1) disturbances directly affect forest productivity (e.g. via a reduction in leaf area, growing stock or resource-use efficiency), and 2) disturbance susceptibility is often coupled to a certain development phase of the forest with productivity determining the time a forest is in this specific phase of susceptibility. The objective of this paper is to provide an overview of forest productivity changes in different forest regions in Europe under climate change, and partition these changes into effects induced by climate change alone and by climate change and disturbances. We present projections of climate change impacts on forest productivity from state-of-the-art forest models that dynamically simulate forest productivity and the effects of the main European disturbance agents (fire, storm, insects), driven by the same climate scenario in seven forest case studies along a large climatic gradient throughout Europe. Our study shows that, in most cases, including disturbances in the simulations exaggerate ongoing productivity declines or cancel out productivity gains in response to climate change. In fewer cases, disturbances also increase productivity or buffer climate-change induced productivity losses, e.g. because low severity fires can alleviate resource competition and increase fertilization. Even though our results cannot simply be extrapolated to other types of forests and disturbances, we argue that it is necessary to interpret climate change-induced productivity and disturbance changes jointly to capture the full range of climate change impacts on forests and to plan adaptation measures.

Responses of eastern Chinese coastal salt marshes to sea-level rise combined with vegetative and sedimentary processes.

The impacts of sea-level rise (SLR) on coastal ecosystems have attracted worldwide attention in relation to global change. In this study, the salt marsh model for the Yangtze Estuary (SMM-YE, developed in China) and the Sea Level Affecting Marshes Model (SLAMM, developed in the U.S.) were used to simulate the effects of SLR on the coastal salt marshes in eastern China. The changes in the dominant species in the plant community were also considered. Predictions based on the SLAMM indicated a trend of habitat degradation up to 2100; total salt marsh habitat area continued to decline (4-16%) based on the low-level scenario, with greater losses (6-25%) predicted under the high-level scenario. The SMM-YE showed that the salt marshes could be resilient to threats of SLR through the processes of accretion of mudflats, vegetation expansion and sediment trapping by plants. This model predicted that salt marsh areas increased (3-6%) under the low-level scenario. The decrease in the total habitat area with the SMM-YE under the high-level scenario was much lower than the SLAMM prediction. Nevertheless, SLR might negatively affect the salt marsh species that are not adapted to prolonged inundation. An adaptive strategy for responding to changes in sediment resources is necessary in the Yangtze Estuary.

Comparison of the effects of symmetric and asymmetric temperature elevation and CO2 enrichment on yield and evapotranspiration of winter wheat (Triticum aestivum L.).

Under the changing climate, asymmetric warming pattern would be more likely during day and night time, instead of symmetric one. Concurrently, the growth responses and water use of plants may be different compared with those estimated based on symmetric warming. In this work, it was compared with the effects of symmetric (ETs) and asymmetric (ETa) elevation of temperature alone, and in interaction with elevated carbon dioxide concentration (EC), on the grain yield (GY) and evapotranspiration in winter wheat (Triticum aestivum L.) based on pot experiment in the North China Plain (NCP). The experiment was carried out in six enclosed-top chambers with following climate treatments: (1) ambient temperature and ambient CO2 (CON), (2) ambient temperature and elevated CO2 (EC), (3) elevated temperature and ambient CO2 (ETs; ETa), and (4) elevated temperature and elevated CO2 (ECETs, ECETa). In symmetric warming, temperature was increased by 3°C and in asymmetric one by 3.5°C during night and 2.5°C during daytime, respectively. As a result, GY was in ETa and ETs 15.6 (P < 0.05) and 10.3% (P < 0.05) lower than that in CON. In ECETs and ECETa treatments, GY was 14.9 (P < 0.05) and 9.1% (P < 0.05) higher than that in CON. Opposite to GY, evapotranspiration was 7.8 (P < 0.05) and 17.9% (P < 0.05) higher in ETa and ETs treatments and 7.2 (P < 0.05) and 2.1% (P > 0.05) lower in ECETs and ECETa treatments compared with CON. Thus, GY of wheat could be expected to increase under the changing climate with concurrent elevation of CO2 and temperature as a result of increased WUE under the elevated CO2. However, the gain would be lower under ETa than that estimated based on ETs due to higher evapotranspiration.

Controls of evapotranspiration and CO2 fluxes from scots pine by surface conductance and abiotic factors.

Evapotranspiration (E) and CO2 flux (Fc ) in the growing season of an unusual dry year were measured continuously over a Scots pine forest in eastern Finland, by eddy covariance techniques. The aims were to gain an understanding of their biological and environmental control processes. As a result, there were obvious diurnal and seasonal changes in E, Fc , surface conductance (gc ), and decoupling coefficient (Ω), showing similar trends to those in radiation (PAR) and vapour pressure deficit (δ). The maximum mean daily values (24-h average) for E, Fc , gc , and Ω were 1.78 mmol m(-2) s(-1), -11.18 µmol m(-2) s(-1), 6.27 mm s(-1), and 0.31, respectively, with seasonal averages of 0.71 mmol m(-2) s(-1), -4.61 µmol m(-2) s(-1), 3.3 mm s(-1), and 0.16. E and Fc were controlled by combined biological and environmental variables. There was curvilinear dependence of E on gc and Fc on gc . Among the environmental variables, PAR was the most important factor having a positive linear relationship to E and curvilinear relationship to Fc , while vapour pressure deficit was the most important environmental factor affecting gc . Water use efficiency was slightly higher in the dry season, with mean monthly values ranging from 6.67 to 7.48 µmol CO2 (mmol H2O)(-1) and a seasonal average of 7.06 µmol CO2 (µmol H2O)(-1). Low Ω and its close positive relationship with gc indicate that evapotranspiration was sensitive to surface conductance. Mid summer drought reduced surface conductance and decoupling coefficient, suggesting a more biotic control of evapotranspiration and a physiological acclimation to dry air. Surface conductance remained low and constant under dry condition, supporting that a constant value of surface constant can be used for modelling transpiration under drought condition.

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland.

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.

Sensitivity of managed boreal forests in Finland to climate change, with implications for adaptive management.

This study investigated the sensitivity of managed boreal forests to climate change, with consequent needs to adapt the management to climate change. Model simulations representing the Finnish territory between 60 and 70 degrees N showed that climate change may substantially change the dynamics of managed boreal forests in northern Europe. This is especially probable at the northern and southern edges of this forest zone. In the north, forest growth may increase, but the special features of northern forests may be diminished. In the south, climate change may create a suboptimal environment for Norway spruce. Dominance of Scots pine may increase on less fertile sites currently occupied by Norway spruce. Birches may compete with Scots pine even in these sites and the dominance of birches may increase. These changes may reduce the total forest growth locally but, over the whole of Finland, total forest growth may increase by 44%, with an increase of 82% in the potential cutting drain. The choice of appropriate species and reduced rotation length may sustain the productivity of forest land under climate change.

Elevated temperature and CO(2) concentration effects on xylem anatomy of Scots pine.

We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross-sectional area, the greatest decrease occurring in the elevated [CO(2)] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO(2)] and that the effects of temperature were confined to the earlywood.

Mechanical stability of trees under static loads.

Wind affects the structure and functioning of a forest ecosystem continuously and may cause significant economic loss in managed forests by reducing the yield of recoverable timber, increasing the cost of unscheduled thinning and clear-cuttings, and creating problems in forestry planning. Furthermore, broken and uprooted trees within the forest are subject to insect attack and may provide a suitable breeding substrate, endangering the remaining trees. Therefore, an improved understanding of the processes behind the occurrence of wind-induced damage is of interest to many forest ecologists, but may also help managers of forest resources to make appropriate management decisions related to risk management. Using fundamental physics, empirical experiments, and mechanistic model-based approaches in interaction, we can study the susceptibility of tree stands to wind damage as affected by the wind and site and tree/stand characteristics and management. Such studies are not possible based on statistical approaches alone, which are not able to define the causal links between tree parameters and susceptibility to wind damage. The aim of this paper is to review the recent work done related to tree-pulling and wind tunnel experiments and mechanistic modeling approaches to increase our understanding of the mechanical stability of trees under static loading.

Annual and seasonal variation of sap flow and conductance of pine trees grown in elevated carbon dioxide and temperature.

Measurements of sap flow, crown structure, and microclimate were used to estimate the transpiration of individual 30-year-old Pinus sylvestris L. trees grown in elevated temperature and CO2. The trees were enclosed in closed-top chambers and exposed either to current ambient conditions (CON), or elevated CO2 (+350 micromol mol(-1); EC), or elevated temperature (+2 to +6 degrees C; ET) or a combination of EC and ET (ECT) since 1996, and the measurements were made from 1999 to 2001. EC significantly increased annual sap flow per tree (Ft.m) by 14% in 1999, but reduced it by 13% in 2000 and 16% in 2001. The CO2-induced increase in Ft.m in 1999 was due to a large increase in foliage area of trees, which more than compensated for a small decrease in crown conductance (Gc). The CO2-induced decreases in Ft.m in 2000 and 2001 resulted from a pronounced decline in Gc, which was much greater than the increase in foliage area. The CO2-induced increase in sensitivity of Gc at high vapour pressure deficit (VPD) did not alter the general response of sap flow to CO2 enrichment, but it did affect the diurnal courses of sap flow on some days during the main growing season (days 150-240). ET increased Ft.m by 53%, 45%, and 57% in 1999, 2000, and 2001, respectively, attributable to the combined effects of greater foliage area and maximum crown conductance, lower stomatal sensitivity to high VPD, and higher transpiration demand relative to the control treatments. There was no significant interaction between CO2 and temperature on sap flow, because ECT entailed approximately similar patterns of sap flow to ET, suggesting that the temperature played a dominate role in the case of ECT under boreal climate conditions.

Scots pine responses to elevated temperature and carbon dioxide concentration: growth and wood properties.

Growth and wood properties of 20-year-old Scots pine (Pinus sylvestris L.) trees were studied for 6 years in 16 closed chambers providing a factorial combination of two temperature regimes (ambient and elevated) and two carbon dioxide concentrations ([CO2]) (ambient and twice ambient). The elevation of temperature corresponded to the predicted effect at the site of a doubling in atmospheric [CO2]. Annual height and radial growth and wood properties were analyzed during 1997-2002. Physical wood properties analyzed included early- and latewood widths and their proportions, intra-ring wood densities, early- and latewood density and mean fiber length. Chemical wood properties analyzed included concentrations of acetone-soluble extractives, lignin, cellulose and hemicellulose. There were no significant treatment effects on height growth during the 6-year study. Elevated [CO2] increased ring width by 66 and 47% at ambient and elevated temperatures, respectively. At ambient [CO2], elevated temperature increased ring width by 19%. Increased ring width in response to elevated [CO2] resulted from increases in both early- and latewood width; however, there was no effect of the treatments on early- and latewood proportions. Mean wood density, earlywood density and fiber length increased in response to elevated temperature. The chemical composition of wood was affected by elevated [CO2], which reduced the cellulose concentration, and by elevated temperature, which reduced the concentration of acetone-soluble extractives. Thus, over the 6-year period, radial growth was significantly increased by elevated [CO2], and some wood properties were significantly affected by elevated temperature or elevated [CO2], or both, indicating that climate change may affect the material properties of wood.

Wood properties of Scots pines (Pinus sylvestris) grown at elevated temperature and carbon dioxide concentration.

Impacts of elevated temperature and carbon dioxide concentration ([CO2]) on wood properties of 15-year-old Scots pines (Pinus sylvestris L.) grown under conditions of low nitrogen supply were investigated in open-top chambers. The treatments consisted of (i) ambient temperature and ambient [CO2] (AT+AC), (ii) ambient temperature and elevated [CO2] (AT+EC), (iii) elevated temperature and ambient [CO2] (ET+AC) and (iv) elevated temperature and elevated [CO2] (ET+EC). Wood properties analyzed for the years 1992-1994 included ring width, early- and latewood width and their proportions, intra-ring wood density (minimum, maximum and mean, as well as early- and latewood densities), mean fiber length and chemical composition of the wood (cellulose, hemicellulose, lignin and acetone extractive concentration). Absolute radial growth over the 3-year period was 54% greater in AT+EC trees and 30 and 25% greater in ET+AC and ET+EC trees, respectively, than in AT+AC trees. Neither elevated temperature nor elevated [CO2] had a statistically significant effect on ring width, early- and latewood widths or their proportions. Both latewood density and maximum intra-ring density were increased by elevated [CO2], whereas fiber length was increased by elevated temperature. Hemicellulose concentration decreased and lignin concentration increased significantly in response to elevated temperature. There were no statistically significant interaction effects of elevated temperature and elevated [CO2] on the wood properties, except on earlywood density.

Diameter growth of Scots pine (Pinus sylvestris) trees grown at elevated temperature and carbon dioxide concentration under boreal conditions.

We investigated the impacts of elevated temperature and carbon dioxide concentration ([CO2]) on diameter growth of Scots pine (Pinus sylvestris L.), aged about 20 years, grown with a low nitrogen supply in closed chambers at (i) ambient temperature and [CO2] (AT+AC), (ii) ambient temperature and elevated [CO2] (AT+EC), (iii) elevated temperature and ambient [CO2] (ET+AC), and (iv). elevated temperature and [CO2] (ET+EC). Each treatment was replicated four times. Diameter growth was monitored with a band dendrograph at 15-min intervals throughout the growing seasons of 1997, 1998 and 1999. Over the monitoring period, diameter growth began 2-3 weeks earlier in trees in the ET+EC and ET+AC chambers than in trees in the AT+AC and AT+EC chambers. However, the cessation of growth occurred about a week later in trees in the ET+EC, ET+AC and AT+EC chambers compared with the AT+AC chambers. The duration of the growing season was 115 and 108 days in the ET+EC and ET+AC chambers, respectively, and 95 and 84 days in the AT+EC and AT+AC chambers, respectively. The ET+AC and ET+EC treatments enhanced diameter growth most early in the growing season, whereas in trees in the AT+AC and AT+EC treatments diameter growth rate was highest in the middle of the growing season. Diameter growth rate leveled off more slowly in trees in the ET+EC and AT+EC treatments than in the other treatments. The growth response to elevated T, elevated [CO2] or both decreased with time and it was less than the maximum observed in other studies for small seedlings and under optimal growth conditions. Nevertheless, cumulative diameter growth for the 3-year period was 67% greater in trees in the ET+EC treatment, and 57 and 26% greater in trees in the AT+EC and ET+AC treatments, respectively, compared with trees in the AT+AC treatment. Over the 3 years, [CO2] had a statistically significant (P < 0.10) effect on both absolute and relative diameter growth, but the interaction between [CO2] and temperature was not significant.