Enhancing boreal forest resilience: A four-year impact of biochar on soil quality and fungal communities.

Boreal forests commonly suffer from nutrient deficiency due to restricted biological activity and decomposition. Biochar has been used as a promising strategy to improve soil quality, yet its impacts on forest soil microbes, particularly in cold environment, remains poorly understood. In this study, we investigated the effects of biochar, produced at different pyrolysis temperatures (500 °C and 650 °C) and applied at different amounts (0.5 kg·m-2 and 1.0 kg·m-2), on soil property, soil enzyme activity, and fungal community dynamics in a boreal forest over a span of two to four years. Our results showed that, four-year post-application of biochar produced at 650 °C and applied at 1.0 kg·m-2, significantly increased the relative abundance of Mortierellomycota and enhanced fungal species richness, α-diversity and evenness compared to the control (CK) (P < 0.05). Notably, the abundance of Phialocephala fortinii increased with the application of biochar produced at 500 °C and applied at 0.5 kg·m-2, exhibiting a positively correlation with the carbon cycling-related enzyme ß-cellobiosidase. Functionally, distinct fungal gene structures were formed between different biochar pyrolysis temperatures, and between application amounts in four-year post-biochar application (P < 0.05). Additionally, correlation analyses revealed the significance of the duration post-biochar application on the soil properties, soil extracellular enzymes, soil fungal dominant phyla, fungal community and gene structures (P < 0.01). The interaction between biochar pyrolysis temperature and application amount significantly influenced fungal α-diversity (P < 0.01). Overall, these findings provide theoretical insights and practical application for biochar as soil amendment in boreal forest ecosystems.

The effect of ectomycorrhizal fungal exposure on nursery-raised Pinus sylvestris seedlings: plant transpiration under short-term drought, root morphology and plant biomass.

Drought is a major environmental stressor that limits seedling growth. Several studies have found that some ectomycorrhizal fungi may increase the drought tolerance of nursery-raised seedlings. However, the precise role that different ectomycorrhizal fungi species play in drought tolerance remains unclear. We evaluated the transpiration rate of Pinus sylvestris seedlings under drought stress in greenhouse conditions by exposing seedlings to 10 ectomycorrhizal fungi species, with different functional traits (exploration type and hydrophobicity), and to 3 natural soil inoculums. We measured the transpiration and water potential of the seedlings during a 10-day drought period and a 14-day recovery period. We then analyzed their root morphology, stem, needle, root biomass and needle chlorophyll fluorescence. We showed that exposing seedlings to ectomycorrhizal fungi or soil inoculum had a positive effect on their transpiration rate during the driest period and through the recovery phase, leading to 2- to 3-fold higher transpiration rates compared with the nonexposed control seedlings. Seedlings exposed to medium-distance ectomycorrhizal fungi performed better than other exploration types under drought conditions, but ectomycorrhizal fungi hydrophobicity did not seem to affect the seedlings response to drought. No significant differences were observed in biomass accumulation and root morphology between the seedlings exposed to different ectomycorrhizal fungi species and the control. Our results highlight the positive and species-specific effect of ectomycorrhizal fungi exposure on drought tolerance in nursery-raised Scots pine seedlings. The studied ectomycorrhizal fungi functional traits may not be sufficient to predict the seedling response to drought stress, thus physiological studies across multiple species are needed to draw the correct conclusion. Our findings have potential practical implications for enhancing seedling drought tolerance in nursery plant production.

High organic carbon content constricts the potential for stable organic carbon accrual in mineral agricultural soils in Finland.

Sequestering carbon into agricultural soils is considered as a means of mitigating climate change. We used agronomic soil test results representing c. 95% of the farmed land area in Finland to estimate the potential of the uppermost 15 cm soil layer of mineral agricultural soils to sequester organic carbon (OC) and to contribute to the mitigation of climate change. The estimation of the maximum capacity of mineral matter to protect OC in stable mineral-associated form was based on the theory that clay and fine-sized (fines = clay + silt) particles have a limited capacity to protect OC. In addition, we used the clay/OC and fines/OC ratios to identify areas with a risk of erosion and reduced productivity, thus indicating priority areas potentially benefitting from the increased soil OC contents. We found that 32-40% of the mineral agricultural soils in Finland have the potential to further accumulate mineral-associated OC (MOC), while in the majority of soils, the current OC stock in the uppermost 15 cm exceeded the capacity of mineral matter to protect OC. The nationwide soil OC sequestration potential of the uppermost 15 cm in mineral agricultural soils ranged between 0.21 and 0.26 Tg, which corresponds to less than 2% of annual greenhouse gas emissions in Finland. The fields with the highest potential for SOC accrual were found in the southern and southwestern parts of the country, including some of the most intensively cultivated high-clay soils. Although the nationwide potential for additional OC sequestration was estimated to be relatively small, the current OC storage in Finnish arable mineral soils (0-15 cm) is large, 128 Tg. Farming practices enabling maximum OC input into the soil play an important role as a tool for mitigating the loss of carbon from high-OC soils in the changing climate. Furthermore, especially in high-clay areas with potential for MOC accrual, efforts to increase soil OC could help improve soil structural stability and therefore reduce erosion and the loss of nutrients to the aquatic environments.

Ligneous amendments increase soil organic carbon content in fine-textured boreal soils and modulate N2O emissions.

Organic soil amendments are used to improve soil quality and mitigate climate change. However, their effects on soil structure, nutrient and water retention as well as greenhouse gas (GHG) emissions are still poorly understood. The purpose of this study was to determine the residual effects of a single field application of four ligneous soil amendments on soil structure and GHG emissions. We conducted a laboratory incubation experiment using soil samples collected from an ongoing soil-amendment field experiment at Qvidja Farm in south-west Finland, two years after a single application of four ligneous biomasses. Specifically, two biochars (willow and spruce) produced via slow pyrolysis, and two mixed pulp sludges from paper industry side-streams were applied at a rate of 9-22 Mg ha-1 mixed in the top 0.1 m soil layer. An unamended fertilized soil was used as a control. The laboratory incubation lasted for 33 days, during which the samples were kept at room temperature (21°C) and at 20%, 40%, 70% or 100% water holding capacity. Carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) fluxes were measured periodically after 1, 5, 12, 20 and 33 days of incubation. The application of ligneous soil amendments increased the pH of the sampled soils by 0.4-0.8 units, whereas the effects on soil organic carbon content and soil structure varied between treatments. The GHG exchange was dominated by CO2 emissions, which were mainly unaffected by the soil amendment treatments. The contribution of soil CH4 exchange was negligible (nearly no emissions) compared to soil CO2 and N2O emissions. The soil N2O emissions exhibited a positive exponential relationship with soil moisture. Overall, the soil amendments reduced N2O emissions on average by 13%, 64%, 28%, and 37%, at the four soil moisture levels, respectively. Furthermore, the variation in N2O emissions between the amendments correlated positively with their liming effect. More specifically, the potential for the pulp sludge treatments to modulate N2O emissions was evident only in response to high water contents. This tendency to modulate N2O emissions was attributed to their capacity to increase soil pH and influence soil processes by persisting in the soil long after their application.

The impact of biochar on wood-inhabiting bacterial community and its function in a boreal pine forest.

Biochar is considered to be a possible means of carbon sequestration to alleviate climate change. However, the dynamics of the microbial community during wood decomposition after biochar application remain poorly understood. In this study, the wood-inhabiting bacterial community composition and its potential functions during a two-year decomposition period after the addition of different amounts of biochar (0.5 kg m-2 and 1.0 kg m-2), and at different biochar pyrolysis temperatures (500 °C and 650 °C), in a boreal Scots pine forest, were analyzed using Illumina NovaSeq sequencing combined with Functional Annotation of Prokaryotic Taxa (FAPROTAX). The results showed that the wood decomposition rates increased after biochar addition to the soil surface in the second year. Treatment with biochar produced at high temperatures increased the diversity of wood-inhabiting bacteria more than that produced at low temperatures (P < 0.05). The wood-inhabiting bacterial diversity and species richness decreased with decomposition time. The biochar treatments changed the wood-inhabiting bacterial community structure during the decomposition period. The pyrolysis temperature and the amount of applied biochar had no effect on the bacterial community structure but shifted the abundance of certain bacterial taxa. Similarly, biochar application shifted the wood-inhabiting bacterial community function in the first year, but not in the second year. The wood-inhabiting bacterial community and function were affected by soil pH, soil water content, and soil total nitrogen. The results provide useful information on biochar application for future forest management practices. Long-term monitoring is needed to better understand the effects of biochar application on nutrient cycling in boreal forests.

Microbial biodiversity contributes to soil carbon release: a case study on fire disturbed boreal forests.

Microbial communities often possess enormous diversity, raising questions about whether this diversity drives ecosystem functioning, especially the influence of diversity on soil decomposition and respiration. Although functional redundancy is widely observed in soil microorganisms, evidence that species occupy distinct metabolic niches has also emerged. In this paper, we found that apart from the environmental variables, increases in microbial diversity, notably bacterial diversity, lead to an increase in soil C emissions. This was demonstrated using structural equation modelling (SEM), linking soil respiration with naturally differing levels of soil physio-chemical properties, vegetation coverage, and microbial diversity after fire disturbance. Our SEMs also revealed that models including bacterial diversity explained more variation of soil CO2 emissions (about 45%) than fungal diversity (about 38%). A possible explanation of this discrepancy is that fungi are more multifunctional than bacteria and, therefore, an increase in fungal diversity does not necessarily change soil respiration. Further analysis on functional gene structure suggested that bacterial and fungal diversities mainly explain the potential decomposition of recalcitrant C compare with that of labile C. Overall, by incorporating microbial diversity and the environmental variables, the predictive power of models on soil C emission was significantly improved, indicating microbial diversity is crucial for predicting ecosystem functions.

Mycelium chemistry differs markedly between ectomycorrhizal and arbuscular mycorrhizal fungi.

The chemical quality of soil carbon (C) inputs is a major factor controlling litter decomposition and soil C dynamics. Mycorrhizal fungi constitute one of the dominant pools of soil microbial C, while their litter quality (chemical proxies of litter decomposability) is understood poorly, leading to major uncertainties in estimating soil C dynamics. We examined litter decomposability of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal species using samples obtained from in vitro cultivation. We showed that the chemical composition of AM and EM fungal mycelium differs significantly: EM fungi have higher concentrations of labile (water-soluble, ethanol-soluble) and recalcitrant (non-extractable) chemical components, while AM fungi have higher concentrations of acid-hydrolysable components. Our results imply that differences in decomposability traits among mycorrhizal fungal guilds represent a critically important driver of the soil C cycle, which could be as vital as is recognized for differences among aboveground plant litter.

Soil organic carbon stocks and dynamics in a mollisol region: A 1980s-2010s study.

Mollisols are globally distributed in grain-producing regions, and soil organic carbon (SOC) dynamics in mollisol regions are closely related to food security. Regional climate, land use and cover, and field management practice have massively changed since the 1980s in mollisol region in Northeast China, however, the dynamics of topsoil and profile SOC stocks and their distribution have not updated. To explore the dynamics of SOC stocks and their horizontal and vertical distributions in the 1980s-2010s, we took the mollisol region in Northeast China as an example location to conduct profile-scale soil surveys. The in situ surveys indicated that the topsoil SOC stock (0-20 cm) remained relatively stable throughout the 1980s, 2000s, and 2010s, and was 57.3 ± 5.5, 58.2 ± 3.3, and 57.4 ± 4.4 t C ha-1, respectively. The average profile SOC stock (1 m) increased from 148.9 ± 18.5 t C ha-1 in the 1980s to 162.0 ± 14.0 t C ha-1 in the 2010s. A slowdown in land reclamation and implementation of conservation tillage helped maintain and restore SOC stocks. Although the overall SOC stock tended to accumulate, the study area suffered an increasingly unbalanced redistribution of SOC related to severe soil erosion. Soil particles and SOC at erosional positions such as backslope were stripped from the soil surface, leading to attenuated soil thickness and SOC stock; SOC-rich sediment accumulated and was buried at depositional positions, especially at the foot-slope, increasing the soil thickness and SOC stock. These results confirmed that not only the total SOC stock, but also changes in SOC spatial distribution deserve great attention. This study provides a platform to examine and modify the simulation effectiveness of carbon-cycling models, as well as solid foundations for optimal global mollisols management.

Soil Fungal Community Structure in Boreal Pine Forests: From Southern to Subarctic Areas of Finland.

The boreal forest environment plays an important role in the global C cycle due to its high carbon storage capacity. However, relatively little is known about the forest fungal community at a regional scale in boreal forests. In the present study, we have re-analyzed the data from our previous studies and highlighted the core fungal community composition and potential functional groups in three forests dominated by Scots pine (Pinus sylvestris L.) in Finland, and identified the fungal generalists that appear across geographic locations despite differences in local conditions. The three forests represent subarctic, northern and southern boreal forest, and are all in an un-managed state without human interference or management. The subarctic and northern areas are subject to reindeer grazing. The results showed that the three locations formed distinct fungal community structures (P < 0.05). Compared to the two northern locations, the southern boreal forest harbored a greater abundance of Zygomycota, Lactarius, Mortierella Umbelopsis, and Tylospora, in which aspect there were no differences between the two northern forests. Cortinarius, Piloderma, and Suillus were the core fungal genera in the boreal Scots pine forest. Functionally, the southern boreal forest harbored a greater abundance of saprotroph, endophytes and fungal parasite-lichen, whereas a greater abundance of ectomycorrhizal fungi was observed in the northern boreal forests. Moreover, the pathotroph and wood saprotrophs were commonly present in these three regions. The three locations formed two distinct fungal community functional structures, by which the southern forest was clearly separated from the two northern forests, suggesting a distance-decay relationship via geographic location. This study provides useful information for better understanding the common fungal communities and functions in boreal forests in different geographical locations.

Root presence modifies the long-term decomposition dynamics of fungal necromass and the associated microbial communities in a boreal forest.

Recent studies have highlighted that dead fungal mycelium represents an important fraction of soil carbon (C) and nitrogen (N) inputs and stocks. Consequently, identifying the microbial communities and the ecological factors that govern the decomposition of fungal necromass will provide critical insight into how fungal organic matter (OM) affects forest soil C and nutrient cycles. Here, we examined the microbial communities colonising fungal necromass during a multiyear decomposition experiment in a boreal forest, which included incubation bags with different mesh sizes to manipulate both plant root and microbial decomposer group access. Necromass-associated bacterial and fungal communities were taxonomically and functionally rich throughout the 30 months of incubation, with increasing abundances of oligotrophic bacteria and root-associated fungi (i.e., ectomycorrhizal, ericoid mycorrhizal and endophytic fungi) in the late stages of decomposition in the mesh bags to which they had access. Necromass-associated ß-glucosidase activity was highest at 6 months, while leucine aminopeptidase peptidase was highest at 18 months. Based on an asymptotic decomposition model, root presence was associated with an initial faster rate of fungal necromass decomposition, but resulted in higher amounts of fungal necromass retained at later sampling times. Collectively, these results indicate that microbial community composition and enzyme activities on decomposing fungal necromass remain dynamic years after initial input, and that roots and their associated fungal symbionts result in the slowing of microbial necromass turnover with time.

Mechanisms of Carbon Sequestration in Highly Organic Ecosystems - Importance of Chemical Ecology.

Organic matter decomposition plays a major role in the cycling of carbon (C) and nutrients in terrestrial ecosystems across the globe. Climate change accelerates the decomposition rate to potentially increase the release of greenhouse gases and further enhance global warming in the future. However, fractions of organic matter vary in turnover times and parts are stabilized in soils for longer time periods (C sequestration). Overall, a better understanding of the mechanisms underlying C sequestration is needed for the development of effective mitigation policies to reduce land-based production of greenhouse gases. Known mechanisms of C sequestration include the recalcitrance of C input, interactions with soil minerals, aggregate formation, as well as its regulation via abiotic factors. In this Minireview, we discuss the mechanisms behind C sequestration including the recently emerging significance of biochemical interactions between organic matter inputs that lead to C stabilization.

Long-term effects of forest fires on soil greenhouse gas emissions and extracellular enzyme activities in a hemiboreal forest.

Fire is the most important natural disturbance in boreal forests, and it has a major role regulating the carbon (C) budget of these systems. With the expected increase in fire frequency, the greenhouse gas (GHG) budget of boreal forest soils may change. In order to understand the long-term nature of the soil-atmosphere GHG exchange after fire, we established a fire chronosequence representing successional stages at 8, 19, 34, 65, 76 and 179 years following stand-replacing fires in hemiboreal Scots pine forests in Estonia. Changes in extracellular activity, litter decomposition, vegetation biomass, and soil physicochemical properties were assessed in relation to carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) emissions. Soil temperature was highest 8 years after fire, whereas soil moisture varied through the fire chronosequences without a consistent pattern. Litter decomposition and CO2 efflux were still lower 8 years after fire compared with pre-fire levels (179 years after fire). Both returned to pre-fire levels before vegetation re-established, and CO2 efflux was only strongly responsive to temperature from 19 years after fire onward. Recovery of CO2 efflux in the long term was associated with a moderate effect of fire on enzyme activity, the input of above- and below-ground litter carbon, and the re-establishment of vegetation. Soil acted as a CH4 sink and N2O source similarly in all successional stages. Compared with soil moisture and time after fire, soil temperature was the most important predictor for both GHGs. The re-establishment of overstorey and vegetation cover (mosses and lichens) might have caused an increase in CH4 and N2O effluxes in the studied areas, respectively.

Restriction of plant roots in boreal forest organic soils affects the microbial community but does not change the dominance from ectomycorrhizal to saprotrophic fungi.

Boreal forest soils store significant amounts of carbon and are cohabited by saprotrophic and ectomycorrhizal fungi (ECM). The 'Gadgil effect' implies antagonistic interactions between saprotrophic fungi and ECM. Plant photosynthates support the competitive fitness of the ECM, and may also shape the soil bacterial communities. Many 'Gadgil effect' experiments have focused on litter layer (OL) or have litter and root-fragments present, and thus possibly favor the saprotrophs. We compared how the restriction of plant roots and exudates affect soil microbial community structures in organic soil (mixed OF and OH). For this, we established a 3-yr field experiment with 3 different mesh treatments affecting the penetration of plant roots and external fungal hyphae. Exclusion of plant photosynthates induced modest changes in both fungal and bacterial community structures, but not to potential functionality of the microbial community. The microbial community was resilient towards rather short-term disturbances. Contrary to the 'Gadgil effect', mesh treatments restricting the entrance of plant roots and external fungal hyphae did not favor saprotrophs that originally inhabited the soil. Thus, we propose that different substrate preferences (fresh litter vs. fermented or humified soil), rather than antagonism, maintain the spatial separation of saprotrophs and mycorrhizal fungi in boreal forest soils.

Decomposition of spruce wood and release of volatile organic compounds depend on decay type, fungal interactions and enzyme production patterns.

Effect of three wood-decaying fungi on decomposition of spruce wood was studied in solid-state cultivation conditions for a period of three months. Two white rot species (Trichaptum abietinum and Phlebia radiata) were challenged by a brown rot species (Fomitopsis pinicola) in varying combinations. Wood decomposition patterns as determined by mass loss, carbon to nitrogen ratio, accumulation of dissolved sugars and release of volatile organic compounds (VOCs) were observed to depend on both fungal combinations and growth time. Similar dependence of fungal species combination, either white or brown rot dominated, was observed for secreted enzyme activities on spruce wood. Fenton chemistry suggesting reduction of Fe3+ to Fe2+ was detected in the presence of F. pinicola, even in co-cultures, together with substantial degradation of wood carbohydrates and accumulation of oxalic acid. Significant correlation was perceived with two enzyme activity patterns (oxidoreductases produced by white rot fungi; hydrolytic enzymes produced by the brown rot fungus) and wood degradation efficiency. Moreover, emission of four signature VOCs clearly grouped the fungal combinations. Our results indicate that fungal decay type, either brown or white rot, determines the loss of wood mass and decomposition of polysaccharides as well as the pattern of VOCs released upon fungal growth on spruce wood.

Plant roots increase both decomposition and stable organic matter formation in boreal forest soil.

Boreal forests are ecosystems with low nitrogen (N) availability that store globally significant amounts of carbon (C), mainly in plant biomass and soil organic matter (SOM). Although crucial for future climate change predictions, the mechanisms controlling boreal C and N pools are not well understood. Here, using a three-year field experiment, we compare SOM decomposition and stabilization in the presence of roots, with exclusion of roots but presence of fungal hyphae and with exclusion of both roots and fungal hyphae. Roots accelerate SOM decomposition compared to the root exclusion treatments, but also promote a different soil N economy with higher concentrations of organic soil N compared to inorganic soil N accompanied with the build-up of stable SOM-N. In contrast, root exclusion leads to an inorganic soil N economy (i.e., high level of inorganic N) with reduced stable SOM-N build-up. Based on our findings, we provide a framework on how plant roots affect SOM decomposition and stabilization.

Ericoid plant species and Pinus sylvestris shape fungal communities in their roots and surrounding soil.

Root-colonizing fungi can form mycorrhizal or endophytic associations with plant roots, the type of association depending on the host. We investigated the differences and similarities of the fungal communities of three boreal ericoid plants and one coniferous tree, and identified the community structure of fungi utilizing photosynthates from the plants studied. The fungal communities of roots and soils of Vaccinium myrtillus, Vaccinium vitis-idaea, Calluna vulgaris and Pinus sylvestris were studied in an 18-month-long experiment where the plants were grown individually in natural substrate. Photosynthates utilizing fungi were detected with DNA stable-isotope probing using 13 CO2 (13 C-DNA-SIP). The results indicated that the plants studied provide different ecological niches preferred by different fungal species. Those fungi which dominated the community in washed roots had also the highest 13 C-uptake. In addition, a common root endophyte without confirmed mycorrhizal status also obtained 13 C from all the plants, indicating close plant-association of this fungal species. We detect several fungal species inhabiting the roots of both ericoid mycorrhizal and ectomycorrhizal plants. Our results highlight that the ecological role of co-occurrence of fungi with different life styles (e.g. mycorrhizal or endophytic) in plant root systems should be further investigated.

Nitrogen balance along a northern boreal forest fire chronosequence.

Fire is a major natural disturbance factor in boreal forests, and the frequency of forest fires is predicted to increase due to climate change. Nitrogen (N) is a key determinant of carbon sequestration in boreal forests because the shortage of N limits tree growth. We studied changes in N pools and fluxes, and the overall N balance across a 155-year non stand-replacing fire chronosequence in sub-arctic Pinus sylvestris forests in Finland. Two years after the fire, total ecosystem N pool was 622 kg ha-1 of which 16% was in the vegetation, 8% in the dead biomass and 76% in the soil. 155 years after the fire, total N pool was 960 kg ha-1, with 27% in the vegetation, 3% in the dead biomass and 69% in the soil. This implies an annual accumulation rate of 2.28 kg ha-1 which was distributed equally between soil and biomass. The observed changes in N pools were consistent with the computed N balance +2.11 kg ha-1 yr-1 over the 155-year post-fire period. Nitrogen deposition was an important component of the N balance. The biological N fixation increased with succession and constituted 9% of the total N input during the 155 post-fire years. N2O fluxes were negligible (≤ 0.01 kg ha-1 yr-1) and did not differ among post-fire age classes. The number and intensity of microbial genes involved in N cycling were lower at the site 60 years after fire compared to the youngest and the oldest sites indicating potential differences in soil N cycling processes. The results suggest that in sub-arctic pine forests, the non-stand-replacing, intermediate-severity fires decrease considerably N pools in biomass but changes in soil and total ecosystem N pools are slight. Current fire-return interval does not seem to pose a great threat to ecosystem productivity and N status in these sub-arctic forests.

Effects of prolonged drought stress on Scots pine seedling carbon allocation.

As the number of drought occurrences has been predicted to increase with increasing temperatures, it is believed that boreal forests will become particularly vulnerable to decreased growth and increased tree mortality caused by the hydraulic failure, carbon starvation and vulnerability to pests following these. Although drought-affected trees are known to have stunted growth, as well as increased allocation of carbon to roots, still not enough is known about the ways in which trees can acclimate to drought. We studied how drought stress affects belowground and aboveground carbon dynamics, as well as nitrogen uptake, in Scots pine (Pinus sylvestris L.) seedlings exposed to prolonged drought. Overall 40 Scots pine seedlings were divided into control and drought treatments over two growing seasons. Seedlings were pulse-labelled with 13CO2 and litter bags containing 15N-labelled root biomass, and these were used to follow nutrient uptake of trees. We determined photosynthesis, biomass distribution, root and rhizosphere respiration, water potential, leaf osmolalities and carbon and nitrogen assimilation patterns in both treatments. The photosynthetic rate of the drought-induced seedlings did not decrease compared to the control group, the maximum leaf specific photosynthetic rate being 0.058 and 0.045 µmol g-1 s-1 for the drought and control treatments, respectively. The effects of drought were, however, observed as lower water potentials, increased osmolalities as well as decreased growth and greater fine root-to-shoot ratio in the drought-treated seedlings. We also observed improved uptake of labelled nitrogen from soil to needles in the drought-treated seedlings. The results indicate acclimation of seedlings to long-term drought by aiming to retain sufficient water uptake with adequate allocation to roots and root-associated mycorrhizal fungi. The plants seem to control water potential with osmolysis, for which sufficient photosynthetic capability is needed.

Ericoid Roots and Mycospheres Govern Plant-Specific Bacterial Communities in Boreal Forest Humus.

In this study, the bacterial populations of roots and mycospheres of the boreal pine forest ericoid plants, heather (Calluna vulgaris), bilberry (Vaccinium myrtillus), and lingonberry (Vaccinium vitis-idaea), were studied by qPCR and next-generation sequencing (NGS). All bacterial communities of mycosphere soils differed from soils uncolonized by mycorrhizal mycelia. Colonization by mycorrhizal hyphae increased the total number of bacterial 16S ribosomal DNA (rDNA) gene copies in the humus but decreased the number of different bacterial operational taxonomic units (OTUs). Nevertheless, ericoid roots and mycospheres supported numerous OTUs not present in uncolonized humus. Bacterial communities in bilberry mycospheres were surprisingly similar to those in pine mycospheres but not to bacterial communities in heather and lingonberry mycospheres. In contrast, bacterial communities of ericoid roots were more similar to each other than to those of pine roots. In all sample types, the relative abundances of bacterial sequences belonging to Alphaproteobacteria and Acidobacteria were higher than the sequences belonging to other classes. Soil samples contained more Actinobacteria, Deltaproteobacteria, Opitutae, and Planctomycetia, whereas Armatimonadia, Betaproteobacteria, Gammaproteobacteria, and Sphingobacteriia were more common to roots. All mycosphere soils and roots harbored bacteria unique to that particular habitat. Our study suggests that the habitation by ericoid plants increases the overall bacterial diversity of boreal forest soils.