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.     

Timing of bud set in Picea abies is regulated by a memory of temperature during zygotic and somatic embryogenesis

It has been shown previously that height growth and bud phenology are influenced by the temperature during zygotic embryogenesis in Picea abies. To test whether this phenomenon operates within individual plants, clones produced through somatic embryogenesis were used. Seeds were from a full-sib family produced in both a cold (outdoor) and a warm (inside a glasshouse) environment. Embryogenic clones derived from mature zygotic embryos from both crossing environments were cultured at 18, 23 and 28 degrees C during the proliferation and embryo maturation steps. After the second growing season in a glasshouse, plants from the warm seed production environment were taller and had significantly later bud set. For the first time, it is also shown that plants are influenced by the in vitro temperature during somatic embryo development. The warmer the temperature, the later the plants formed terminal buds. The differences were similar to those produced by a provenance separation of 4-6 degrees of latitude. The results indicate that there exists a mechanism in P. abies that operates during embryo development and adjusts the timing of bud set in accordance with the temperature conditions in which the mother tree lives. This in turn counteracts negative effects of gene flow among populations located along altitudinal and latitudinal gradients.     

The rate of protein synthesis in needles of Norway spruce (Picea abies): Light stimulation, regulation through photophosphorylation, stress enhancement

The incorporation of ¹⁴C-leucine into the total-protein fraction of needles of Norway spruce (Picea abies [L.] Karst.) during short time incubation was used as a measure of protein synthesis in the light and in the dark. Light saturation curves, obtained for needles of different ages (new flush and 1 and 2 years old) or at different seasons (summer-winter) followed the Michaelis-Menten algorithm, exhibiting marked differences with regard to light saturation (V_max) and the half-saturation constant (K_{5. 2}). The light saturation curves of ATP level (mg g^?1 fresh weight) and of leucine incorporation into protein (nmol mg^?1 h^?1) matched each other, suggesting that photophosphorylation may be decisive for the rate of protein synthesis in the light. This is confirmed by the action spectrum of leucine incorporation. which resembled an action spectrum of leaf photosynthesis, and also by partial inhibition of protein synthesis by 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), an inhibitor of non-cyclic photophosphorylation. Light stimulated protein synthesis showed pronounced seasonal fluctuations with a summer maximum. Furmigation of 5 years old spruce trees for 3 months with SO₂ in combination with O₃ and/or NO₂ caused a distinct enhancement of the protein synthesis rate in the light and, at a reduced absolute level, also in the dark. A similar result was obtained for 40 to 70 years old spruce stands when healthy and sick trees were compared: the latter being afflicted by the novel type of forest decline, which is characterized by yellowish bronze discolouration of sun-exposed older needles and partial loss of older needle generations (3 to 4 years old). The 1 year old needles of the unhealthy trees showed a markedly increased ¹⁴C-leucine incorporation rate which, in the dark, was even more pronounced than in the light. Stress-physiological mechanisms, which could possibly explain this stimulation, are discussed.     

Temperature-respiration relationships differ in mycorrhizal and non-mycorrhizal root systems of Picea abies (L.) Karst

Root respiration has been shown to increase with temperature, but less is known about how this relationship is affected by the fungal partner in mycorrhizal root systems. In order to test respiratory temperature dependence, in particular Q (10) of mycorrhizal and non-mycorrhizal root systems, seedlings of PICEA ABIES (L.) Karst. (Norway spruce) were inoculated with the ectomycorrhizal fungus PILODERMA CROCEUM (Eriksson and Hjortstam, SR430; synonym: PILODERMA FALLAX: [Libert] Stalpers) and planted in soil respiration cuvettes (mycocosms). Temperature dependence of hyphal respiration in sterile cultures was determined and compared with respiration of mycorrhizal roots. Respiration rates of mycorrhizal and non-mycorrhizal root systems as well as sterile cultures were sensitive to temperature. Q (10) of mycorrhizal root systems of 3.0 +/- 0.1 was significantly higher than that of non-mycorrhizal systems (2.5 +/- 0.2). Q (10) of P. CROCEUM in sterile cultures (older than 2 months) was similar to that of mycorrhizal root systems, suggesting that mycorrhizae may have a large influence on the temperature sensitivity of roots in spite of their small biomass. Our results stress the importance of considering mycorrhization when modeling the temperature sensitivity of spruce roots.     

间伐对川西亚高山粗枝云杉人工林细根生物量及碳储量的影响

以川西亚高山50年生粗枝云杉(Picea asperata)人工林为研究对象, 探讨了间伐对粗枝云杉人工林1-5级细根生物量及碳储量的影响。结果表明: 粗枝云杉人工林细根生物量和碳储量随根序等级的增加而显著增加(p < 0.05), 5级根序中1级根生物量及碳储量最小, 5级根生物量及碳储量最大。与对照(间伐0%)相比, 间伐对粗枝云杉人工林林分细根生物量及碳储量有显著影响(p < 0.05); 而对单株细根生物量影响不一, 间伐10%和20%与对照没有显著性差异(p > 0.05)。间伐显著影响生物量在各根序中的分配, 随着间伐强度的增加, 1、2级细根中生物量分配比例增加, 1级细根的生物量增加幅度最大; 3-5级细根的生物量分配比例减小, 5级细根减少幅度最大。其中, 间伐50%显著减少了细根在下层(20-40 cm)土壤中的生物量比例(p < 0.05), 但与间伐20%和30%无显著差异(p > 0.05)。     

Net aboveground biomass declines of four major forest types with forest ageing and climate change in western Canada's boreal forests

Biomass change of the world's forests is critical to the global carbon cycle. Despite storing nearly half of global forest carbon, the boreal biome of diverse forest types and ages is a poorly understood component of the carbon cycle. Using data from 871 permanent plots in the western boreal forest of Canada, we examined net annual aboveground biomass change (ΔAGB) of four major forest types between 1958 and 2011. We found that ΔAGB was higher for deciduous broadleaf (DEC) (1.44 Mg ha^?1 year^?1, 95% Bayesian confidence interval (CI), 1.22–1.68) and early‐successional coniferous forests (ESC) (1.42, CI, 1.30–1.56) than mixed forests (MIX) (0.80, CI, 0.50–1.11) and late‐successional coniferous (LSC) forests (0.62, CI, 0.39–0.88). ΔAGB declined with forest age as well as calendar year. After accounting for the effects of forest age, ΔAGB declined by 0.035, 0.021, 0.032 and 0.069 Mg ha^?1 year^?1 per calendar year in DEC, ESC, MIX and LSC forests, respectively. The ΔAGB declines resulted from increased tree mortality and reduced growth in all forest types except DEC, in which a large biomass loss from mortality was accompanied with a small increase in growth. With every degree of annual temperature increase, ΔAGB decreased by 1.00, 0.20, 0.55 and 1.07 Mg ha^?1 year^?1 in DEC, ESC, MIX and LSC forests, respectively. With every cm decrease of annual climatic moisture availability, ΔAGB decreased 0.030, 0.045 and 0.17 Mg ha^?1 year^?1 in ESC, MIX and LSC forests, but changed little in DEC forests. Our results suggest that persistent warming and decreasing water availability have profound negative effects on forest biomass in the boreal forests of western Canada. Furthermore, our results indicate that forest responses to climate change are strongly dependent on forest composition with late‐successional coniferous forests being most vulnerable to climate changes in terms of aboveground biomass.     

Fine root foraging strategies in Norway spruce forests across a European climate gradient

Fine root acclimation to different environmental conditions is crucial for growth and sustainability of forest trees. Relatively small changes in fine root standing biomass (FRB), morphology or mycorrhizal symbiosis may result in a large change in forest carbon, nutrient and water cycles. We elucidated the changes in fine root traits and associated ectomycorrhizal (EcM) fungi in 12 Norway spruce stands across a climatic and N deposition gradient from subarctic‐boreal to temperate regions in Europe (68°N–48°N). We analysed the standing FRB and the ectomycorrhizal root tip biomass (EcMB, g m^?2) simultaneously with measurements of the EcM root morphological traits (e.g. mean root length, root tissue density (RTD), N% in EcM roots) and frequency of dominating EcM fungi in different stands in relation to climate, soil and site characteristics. Latitude and N deposition explained the greatest proportion of variation in fine root traits. EcMB per stand basal area (BA) increased exponentially with latitude: by about 12.7 kg m^?2 with an increase of 10° latitude from southern Germany to Estonia and southern Finland and by about 44.7 kg m^?2 with next latitudinal 10° from southern to northern Finland. Boreal Norway spruce forests had 4.5 to 11 times more EcM root tips per stand BA, and the tips were 2.1 times longer, with 1.5 times higher RTD and about 1/3 lower N concentration. There was 19% higher proportion of root tips colonized by long‐distance exploration type forming EcM fungi in the southern forests indicating importance of EcM symbiont foraging strategy in fine root nutrient acquisition. In the boreal zone, we predict ca. 50% decrease in EcMB per stand BA with an increase of 2 °C annual mean temperature. Different fine root foraging strategies in boreal and temperate forests highlight the importance of complex studies on respective regulatory mechanisms in changing climate.     

祁连山青海云杉林生物量和碳储量空间分布特征

根据野外调查资料、祁连山地区青海云杉林相图和气象资料,在GIS技术的支持下估算了祁连山地区青海云杉林的生物量和碳储量及其空间分布.结果表明：2008年,研究区青海云杉林平均生物量为209.24 t·hm^-2,总生物量为3.4×10⁷ t;研究区水热条件的差异使青海云杉生物量在地理空间上存在较大的差异性;经度每增加1°,青海云杉生物量增加3.12t·hm^-2;纬度每增加1°,生物量减少3.8 t·hm^-2;海拔每升高100 m,生物量减少0.05 t·hm^-2;2008年,研究区青海云杉林碳密度在70.4～131.1 t·hm^-2,平均碳密度为109.8 t·hm^-2,幼龄林、中龄林、近熟林、成熟林和过熟林的平均碳密度分别为83.8、109.6、122、124.2和117.1 t·hm^-2,研究区青海云杉林总碳储量为1.8×10⁷ t.     

Effects of climate change and management on net climate impacts of production and utilization of energy biomass in Norway spruce with stable age‐class distribution

We studied the effects of climate change and forest management scenarios on net climate impacts (radiative forcing) of production and utilization of energy biomass, in a Norway spruce forest area over an 80‐year simulation period in Finnish boreal conditions. A stable age‐class distribution was used in model‐based analyses to identify purely the management effects under the current and changing climate (SRES B1 and A2 scenarios). The radiative forcing was calculated based on an integrated use of forest ecosystem model simulations and a life cycle assessment (LCA) tool. In this work, forest‐based energy was used to substitute coal, and current forest management (baseline management) was used as a reference management. In alternative management scenarios, the stocking was maintained 20% higher in thinning compared to the baseline management, and nitrogen fertilization was applied. Intensity of energy biomass harvest (e.g. logging residues, coarse roots and stumps) was varied in the final felling of the stands at the age of 80 years. Also, the economic profitability (NPV, 3% interest rate) of integrated production of timber and energy biomass was calculated for each management scenario. Our results showed that compared to the baseline management, climate benefits could be increased by maintaining higher stocking in thinning over rotation, using nitrogen fertilization and harvesting logging residues, stumps and coarse roots in the final felling. Under the gradually changing climate (in both SRES B1 and A2), the climate benefits were lower compared to the current climate. Trade‐offs between NPV and net climate impacts also existed.     

Aboveground carbon sequestration in dry temperate forests varies with climate not fire regime

The storage of carbon in plant tissues and debris has been proposed as a method to offset anthropogenic increases in atmospheric [CO₂]. Temperate forests represent significant above‐ground carbon (AGC) “sinks” because their relatively fast growth and slow decay rates optimise carbon assimilation. Fire is a common disturbance event in temperate forests globally that should strongly influence AGC because: discrete fires consume above‐ground biomass releasing carbon to the atmosphere, and the long‐term application of different fire‐regimes select for specific plant communities that sequester carbon at different rates. We investigated the latter process by quantifying AGC storage at 104 sites in the Sydney Basin Bioregion, Australia, relative to differences in components of the fire regime: frequency, severity and interfire interval. To predict the potential impacts of future climate change on fire/AGC interactions, we stratified our field sites across gradients of mean annual temperature and precipitation and quantified within‐ and between‐factor interactions between the fire and climate variables. In agreement with previous studies, large trees were the primary AGC sink, accounting for ~70% of carbon at sites. Generalised additive models showed that mean annual temperature was the strongest predictor of AGC storage, with a 54% near‐linear decrease predicted across the 6.1°C temperature range experienced at sites. Mean annual precipitation, fire frequency, fire severity and interfire interval were consistently poor predictors of total above‐ground storage, although there were some significant relationships with component stocks. Our results show resilience of AGC to frequent and severe wildfire and suggest temperature mediated decreases in forest carbon storage under future climate change predictions.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Reproduction and seedling establishment of Picea glauca across the northernmost forest‐tundra region in Canada

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest‐tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest‐tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest‐tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest‐tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest‐tundra ecotone.     

Radial growth response of four dominant boreal tree species to climate along a latitudinal gradient in the eastern Canadian boreal forest

To address the central question of how climate change influences tree growth within the context of global warming, we used dendroclimatological analysis to understand the reactions of four major boreal tree species –Populus tremuloides, Betula papyrifera, Picea mariana, and Pinus banksiana– to climatic variations along a broad latitudinal gradient from 46 to 54°N in the eastern Canadian boreal forest. Tree‐ring chronologies from 34 forested stands distributed at a 1° interval were built, transformed into principal components (PCs), and analyzed through bootstrapped correlation analysis over the period 1950–2003 to identify climate factors limiting the radial growth and the detailed radial growth–climate association along the gradient. All species taken together, previous summer temperature (negative influences), and current January and March–April temperatures (positive influences) showed the most consistent relationships with radial growth across the gradient. Combined with the identified species/site‐specific climate factors, our study suggested that moisture conditions during the year before radial growth played a dominant role in positively regulating P. tremuloides growth, whereas January temperature and growing season moisture conditions positively impacted growth of B. papyrifera. Both P. mariana and P. banksiana were positively affected by the current‐year winter and spring or whole growing season temperatures over the entire range of our corridor. Owing to the impacts of different climate factors on growth, these boreal species showed inconsistent responsiveness to recent warming at the transition zone, where B. papyrifera, P. mariana, and P. banksiana would be the most responsive species, whereas P. tremuloides might be the least. Under continued warming, B. papyrifera stands located north of 49°N, P. tremuloides at northern latitudes, and P. mariana and P. banksiana stands located north of 47°N might benefit from warming winter and spring temperatures to enhance their radial growth in the coming decades, whereas other southern stands might be decreasing in radial growth.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

The climate,the fuel and the land use: Long‐term regional variability of biomass burning in boreal forests

The influence of different drivers on changes in North American and European boreal forests biomass burning (BB) during the Holocene was investigated based on the following hypotheses: land use was important only in the southernmost regions, while elsewhere climate was the main driver modulated by changes in fuel type. BB was reconstructed by means of 88 sedimentary charcoal records divided into six different site clusters. A statistical approach was used to explore the relative contribution of (a) pollen‐based mean July/summer temperature and mean annual precipitation reconstructions, (b) an independent model‐based scenario of past land use (LU), and (c) pollen‐based reconstructions of plant functional types (PFTs) on BB. Our hypotheses were tested with: (a) a west‐east northern boreal sector with changing climatic conditions and a homogeneous vegetation, and (b) a north‐south European boreal sector characterized by gradual variation in both climate and vegetation composition. The processes driving BB in boreal forests varied from one region to another during the Holocene. However, general trends in boreal biomass burning were primarily controlled by changes in climate (mean annual precipitation in Alaska, northern Quebec, and northern Fennoscandia, and mean July/summer temperature in central Canada and central Fennoscandia) and, secondarily, by fuel composition (BB positively correlated with the presence of boreal needleleaf evergreen trees in Alaska and in central and southern Fennoscandia). Land use played only a marginal role. A modification towards less flammable tree species (by promoting deciduous stands over fire‐prone conifers) could contribute to reduce circumboreal wildfire risk in future warmer periods.