Soil mycobiomes in native European aspen forests and hybrid aspen plantations have a similar fungal richness but different compositions, mainly driven by edaphic and floristic factors.

Background: The cultivation of short-rotation tree species on non-forest land is increasing due to the growing demand for woody biomass for the future bioeconomy and to mitigate climate change impacts. However, forest plantations are often seen as a trade-off between climate benefits and low biodiversity. The diversity and composition of soil fungal biota in plantations of hybrid aspen, one of the most planted tree species for short-rotation forestry in Northern Europe, are poorly studied. Methods: The goal of this study was to obtain baseline knowledge about the soil fungal biota and the edaphic, floristic and management factors that drive fungal richness and communities in 18-year-old hybrid aspen plantations on former agricultural soils and compare the fungal biota with those of European aspen stands on native forest land in a 130-year chronosequence. Sites were categorized as hybrid aspen (17-18-year-old plantations) and native aspen stands of three age classes (8-29, 30-55, and 65-131-year-old stands). High-throughput sequencing was applied to soil samples to investigate fungal diversity and assemblages. Results: Native aspen forests showed a higher ectomycorrhizal (EcM) fungal OTU richness than plantations, regardless of forest age. Short-distance type EcM genera dominated in both plantations and forests. The richness of saprotrophic fungi was similar between native forest and plantation sites and was highest in the middle-aged class (30-55-year-old stands) in the native aspen stands. The fungal communities of native forests and plantations were significantly different. Community composition varied more, and the natural forest sites were more diverse than the relatively homogeneous plantations. Soil pH was the best explanatory variable to describe soil fungal communities in hybrid aspen stands. Soil fungal community composition did not show any clear patterns between the age classes of native aspen stands. Conclusion: We conclude that edaphic factors are more important in describing fungal communities in both native aspen forest sites and hybrid aspen plantation sites than forest thinning, age, or former land use for plantations. Although first-generation hybrid aspen plantations and native forests are similar in overall fungal diversity, their taxonomic and functional composition is strikingly different. Therefore, hybrid aspen plantations can be used to reduce felling pressure on native forests; however, our knowledge is still insufficient to conclude that plantations could replace native aspen forests from the soil biodiversity perspective.

Growth ranking of hybrid aspen genotypes and its linkage to leaf gas exchange.

BACKGROUND: Afforestation of non-forestland is a new measure by the European Union to enhance climate mitigation and biodiversity. Hybrid aspen (Populus tremula L. × P. tremuloides Michx.) is among the suitable tree species for afforestation to produce woody biomass. However, the best performing genotypic material for intensive biomass production and its physiological adaptation capacity is still unclear. We compared 22 hybrid aspen genotypes growth and leaf physiological characteristics (stomatal conductance, net photosynthesis, intrinsic water-use efficiency) according to their geographical north- or southward transfer (European P. tremula parent from 51° to 60° N and North American P. tremuloides parent from 45° to 54° N) to hemiboreal Estonia (58° N) in a completely randomized design progeny trial. We tested whether the growth ranking of genotypes of different geographical origin has changed from young (3-year-old) to mid-rotation age (13-year-old). The gas exchange parameters were measured in excised shoots in 2021 summer, which was characterised with warmer (+ 4 °C) and drier (17% precipitation from normal) June and July than the long-term average. RESULTS: We found that the northward transfer of hybrid aspen genotypes resulted in a significant gain in growth (two-fold greater diameter at breast height) in comparison with the southward transfer. The early selection of genotypes was generally in good accordance with the middle-aged genotype ranking, while some of the northward transferred genotypes showed improved growth at the middle-age period in comparison with their ranking during the early phase. The genotypes of southward transfer demonstrated higher stomatal conductance, which resulted in higher net photosynthesis, and lower intrinsic water-use efficiency (iWUE) compared with northward transfer genotypes. However, higher photosynthesis did not translate into higher growth rate. The higher physiological activity of southern transferred genotypes was likely related to a better water supply of smaller and consequently more shaded trees under drought. Leaf nitrogen concentration did not have any significant relation with tree growth. CONCLUSIONS: We conclude that the final selection of hybrid aspen genotypes for commercial use should be done in 10-15 years after planting. Physiological traits acquired during periods of droughty conditions may not fully capture the growth potential. Nonetheless, we advocate for a broader integration of physiological measurements alongside traditional traits (such as height and diameter) in genotype field testing to facilitate the selection of climate-adapted planting material for resilient forests.

Phytoremediation capacity of hybrid aspen at sites affected by industry and agriculture.

Fast-growing Populus spp. are well-acknowledged to restore contaminated soils from heavy metals in industrial areas. Thus far, there is no knowledge about the phytoremediation capacity of Populus spp. plantations in hemiboreal Estonia conditions to restore industrially polluted areas. The objective of this study was to assess the soil contamination rate of heavy metals (As, Cd, Co, Cr, Cu, Fe, Mn, Mo, Ni, Pb and Zn) and their uptake by mature hybrid aspen (Populus tremula × Populus tremuloides Michx.) in plantations in different industrial pollution areas (e.g. cement factory, oil shale mining). For the reference, industrially polluted plantations were compared with the low pollution area hybrid aspen plantation on former agricultural soil, which was influenced by fertilization and liming before afforestation. Twenty-one years after afforestation, soil samples were collected from the 0-10 cm topsoil layer. Aboveground biomass sampling was performed for bark and stem wood by ingrowth cores to separate wood formed during early (1-10 years) and late (11-21 years) stand development. Two decades after the afforestation of industrially polluted areas, the heavy metal concentrations in the soil were higher than the reference plantation and the standard reference for unpolluted soils in most cases. The highest concentrations of heavy metals in woody biomass were in the oil shale quarry spoil; because of poor growth, the accumulated pools in aboveground biomass were low. Cd differed from other metals and accumulated less in wood and more in bark. The concentration of heavy metals (Cd, Cr, Cu, Fe, Mn, Ni and Zn) was higher in the first decade of stand formation (1-10 years) than in the last 10 years (11-21 years). High pools of heavy metals were accumulated in aboveground biomass in the reference plantation, indicating the considerable removal of heavy metal residues from the previous fertilization and liming source with harvest. Two decades of afforestation with hybrid aspen is too short for complete ecosystem restoration from heavy metals in industrially polluted areas.

Regional-Scale In-Depth Analysis of Soil Fungal Diversity Reveals Strong pH and Plant Species Effects in Northern Europe.

Soil microbiome has a pivotal role in ecosystem functioning, yet little is known about its build-up from local to regional scales. In a multi-year regional-scale survey involving 1251 plots and long-read third-generation sequencing, we found that soil pH has the strongest effect on the diversity of fungi and its multiple taxonomic and functional groups. The pH effects were typically unimodal, usually both direct and indirect through tree species, soil nutrients or mold abundance. Individual tree species, particularly Pinus sylvestris, Picea abies, and Populus x wettsteinii, and overall ectomycorrhizal plant proportion had relatively stronger effects on the diversity of biotrophic fungi than saprotrophic fungi. We found strong temporal sampling and investigator biases for the abundance of molds, but generally all spatial, temporal and microclimatic effects were weak. Richness of fungi and several functional groups was highest in woodlands and around ruins of buildings but lowest in bogs, with marked group-specific trends. In contrast to our expectations, diversity of soil fungi tended to be higher in forest island habitats potentially due to the edge effect, but fungal richness declined with island distance and in response to forest fragmentation. Virgin forests supported somewhat higher fungal diversity than old non-pristine forests, but there were no differences in richness between natural and anthropogenic habitats such as parks and coppiced gardens. Diversity of most fungal groups suffered from management of seminatural woodlands and parks and thinning of forests, but especially for forests the results depended on fungal group and time since partial harvesting. We conclude that the positive effects of tree diversity on overall fungal richness represent a combined niche effect of soil properties and intimate associations.

The competitive status of trees determines their responsiveness to increasing atmospheric humidity - a climate trend predicted for northern latitudes.

The interactive effects of climate variables and tree-tree competition are still insufficiently understood drivers of forest response to global climate change. Precipitation and air humidity are predicted to rise concurrently at high latitudes of the Northern Hemisphere. We investigated whether the growth response of deciduous trees to elevated air humidity varies with their competitive status. The study was conducted in seed-originated silver birch and monoclonal hybrid aspen stands grown at the free air humidity manipulation (FAHM) experimental site in Estonia, in which manipulated stands (n = 3 for both species) are exposed to artificially elevated relative air humidity (6-7% over the ambient level). The study period included three growing seasons during which the stands had reached the competitive stage (trees were 7 years old in the final year). A significant 'treatment×competitive status' interactive effect on growth was detected in all years in birch (P < 0.01) and in one year in aspen stands (P = 0.015). Competitively advantaged trees were always more strongly affected by elevated humidity. Initially the growth of advantaged and neutral trees of both species remained significantly suppressed in humidified stands. In the following years, dominance and elevated humidity had a synergistic positive effect on the growth of birches. Aspens with different competitive status recovered more uniformly, attaining similar relative growth rates in manipulated and control stands, but preserved a significantly lower total growth yield due to severe initial growth stress. Disadvantaged trees of both species were never significantly affected by elevated humidity. Our results suggest that air humidity affects trees indirectly depending on their social status. Therefore, the response of northern temperate and boreal forests to a more humid climate in future will likely be modified by competitive relationships among trees, which may potentially affect species composition and cause a need to change forestry practices.

Climate change at northern latitudes: rising atmospheric humidity decreases transpiration, N-uptake and growth rate of hybrid aspen.

At northern latitudes a rise in atmospheric humidity and precipitation is predicted as a consequence of global climate change. We studied several growth and functional traits of hybrid aspen (Populus tremula L.×P. tremuloides Michx.) in response to elevated atmospheric humidity (on average 7% over the ambient level) in a free air experimental facility during three growing seasons (2008-2010) in Estonia, which represents northern temperate climate (boreo-nemoral zone). Data were collected from three humidified (H) and three control (C) plots, and analysed using nested linear models. Elevated air humidity significantly reduced height, stem diameter and stem volume increments and transpiration of the trees whereas these effects remained highly significant also after considering the side effects from soil-related confounders within the 2.7 ha study area. Tree leaves were smaller, lighter and had lower leaf mass per area (LMA) in H plots. The magnitude and significance of the humidity treatment effect--inhibition of above-ground growth rate--was more pronounced in larger trees. The lower growth rate in the humidified plots can be partly explained by a decrease in transpiration-driven mass flow of NO(3) (-) in soil, resulting in a significant reduction in the measured uptake of N to foliage in the H plots. The results suggest that the potential growth improvement of fast-growing trees like aspens, due to increasing temperature and atmospheric CO(2) concentration, might be smaller than expected at high latitudes if a rise in atmospheric humidity simultaneously takes place.

Partitioning of carbohydrates and biomass of needles in Scots pine canopy.

The study was aimed at the quantitative evaluation of the temporal and spatial partitioning of non-structural carbohydrates and needle biomass in a canopy of Scots pine (Pinus sylvestris L.) growing in a Myrtillus site type forest stand (predominant in Estonia). The tree canopy was divided into ten equal layers and the material for the spatial partitioning of the investigated characteristics was sampled from all layers. Our findings revealed a significant variation in morphology and in the partitioning of carbohydrates in needles in different layers of the canopy. The study of the temporal dynamics of carbohydrates showed that starch content in needles started to increase in early spring before budbreak, which was accompanied by a decline in soluble carbohydrates. In October, the starch content of needles was low, but the concentration of soluble sugars started to increase attaining a maximum in winter. Regression analysis indicated that before budbreak, the partitioning of soluble sugars in different canopy layers was relatively weakly correlated with the height of the layer; however, a strong correlation was observed for starch. In autumn, when the growth of trees stopped and daily temperatures decreased, the allocation of soluble sugars was correlated with the height of the canopy layer.