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Most trees form symbioses with ectomycorrhizal fungi (EMF) which influence access to growth-limiting soil resources. Mesocosm experiments repeatedly show that EMF species differentially affect plant development, yet whether these effects ripple up to influence the growth of entire forests remains unknown. Here we tested the effects of EMF composition and functional genes relative to variation in well-known drivers of tree growth by combining paired molecular EMF surveys with high-resolution forest inventory data across 15 European countries. We show that EMF composition was linked to a three-fold difference in tree growth rate even when controlling for the primary abiotic drivers of tree growth. Fast tree growth was associated with EMF communities harboring high inorganic but low organic nitrogen acquisition gene proportions and EMF which form contact versus medium-distance fringe exploration types. These findings suggest that EMF composition is a strong bio-indicator of underlying drivers of tree growth and/or that variation of forest EMF communities causes differences in tree growth. While it may be too early to assign causality or directionality, our study is one of the first to link fine-scale variation within a key component of the forest microbiome to ecosystem functioning at a continental scale.

Explaining the large-scale diversity of soil organisms that drive biogeochemical processes—and their responses to environmental change—is critical. However, identifying consistent drivers of belowground diversity and abundance for some soil organisms at large spatial scales remains problematic. Here we investigate a major guild, the ectomycorrhizal fungi, across European forests at a spatial scale and resolution that is—to our knowledge—unprecedented, to explore key biotic and abiotic predictors of ectomycorrhizal diversity and to identify dominant responses and thresholds for change across complex environmental gradients. We show the effect of 38 host, environment, climate and geographical variables on ectomycorrhizal diversity, and define thresholds of community change for key variables. We quantify host specificity and reveal plasticity in functional traits involved in soil foraging across gradients. We conclude that environmental and host factors explain most of the variation in ectomycorrhizal diversity, that the environmental thresholds used as major ecosystem assessment tools need adjustment and that the importance of belowground specificity and plasticity has previously been underappreciated. Analyses of data from 137 forest plots across 20 European countries show that ectomycorrhizal fungal diversity is strongly influenced by environmental and host species factors and provide thresholds to inform ecosystem assessment tools

Forest soils harbor hyper-diverse microbial communities which fundamentally regulate carbon and nutrient cycling across the globe. Directly testing hypotheses on how microbiome diversity is linked to forest carbon storage has been difficult, due to a lack of paired data on microbiome diversity and in situ observations of forest carbon accumulation and storage. Here, we investigated the relationship between soil microbiomes and forest carbon across 238 forest inventory plots spanning 15 European countries. We show that the composition and diversity of fungal, but not bacterial, species is tightly coupled to both forest biotic conditions and a seven-fold variation in tree growth rates and biomass carbon stocks when controlling for the effects of dominant tree type, climate, and other environmental factors. This linkage is particularly strong for symbiotic endophytic and ectomycorrhizal fungi known to directly facilitate tree growth. Since tree growth rates in this system are closely and positively correlated with belowground soil carbon stocks, we conclude that fungal composition is a strong predictor of overall forest carbon storage across the European continent. Soil microbial diversity and composition is thought to play a major role in elemental cycling. Here, the authors analyse a large dataset of soil microbiome and carbon data from European forests and find that soil fungal community composition is a strong predictor of carbon storage.

The northern regions are experiencing considerable changes in winter climate leading to more frequent warm periods, rain-on-snow events and reduced snow pack diminishing the insulation properties of snow cover and increasing soil frost and freeze-thaw cycles. In this study, we investigated how the lack of snow cover, formation of ice encasement and snow compaction affect the size, structure and activities of soil bacterial and fungal communities. Contrary to our hypotheses, snow manipulation treatments over one winter had limited influence on microbial community structure, bacterial or fungal copy numbers or enzyme activities. However, microbial community structure and activities shifted seasonally among soils sampled before snow melt, in early and late growing season and seemed driven by substrate availability. Bacterial and fungal communities were dominated by stress-resistant taxa such as the orders Acidobacteriales, Chaetothyriales and Helotiales that are likely adapted to adverse winter conditions. This study indicated that microbial communities in acidic northern boreal forest soil may be insensitive to direct effects of changing snow cover. However, in long term, the detrimental effects of increased ice and frost to plant roots may alter plant derived carbon and nutrient pools to the soil likely leading to stronger microbial responses.

In northern boreal forests the warming winter climate leads to more frequent snowmelt, rain-on-snow events and freeze-thaw cycles. This may be harmful or even lethal for tree seedlings that spend even a half of the year under snow. We conducted a snow cover manipulation experiment in a natural forest to find out how changing snow conditions affect young Scots pine ( Pinus sylvestris L.) seedlings. The ice encasement (IE), absence of snow (NoSNOW) and snow compaction (COMP) treatments affected ground level temperature, ground frost and subnivean gas concentrations compared to the ambient snow cover (AMB) and led to the increased physical damage and mortality of seedlings. The expression responses of 28 genes related to circadian clock, aerobic and anaerobic energy metabolism, carbohydrate metabolism and stress protection revealed that seedlings were exposed to different stresses in a complex way depending on the thickness and quality of the snow cover. The IE treatment caused hypoxic stress and probably affected roots which resulted in reduced water uptake in the beginning of the growing season. Without protective snowpack in NoSNOW seedlings suffered from cold and drought stresses. The combination of hypoxic and cold stresses in COMP evoked unique transcriptional responses including oxidative stress. Snow cover manipulation induced changes in the expression of several circadian clock related genes suggested that photoreceptors and the circadian clock system play an essential role in the adaptation of Scots pine seedlings to stresses under different snow conditions. Our findings show that warming winter climate alters snow conditions and consequently causes Scots pine seedlings various abiotic stresses, whose effects extend from overwintering to the following growing season.

At high latitudes, the climate has warmed at twice the rate of the global average with most changes observed in autumn, winter and spring. Increasing winter temperatures and wide temperature fluctuations are leading to more frequent rain-on-snow events and freeze-thaw cycles causing snow compaction and formation of ice layers in the snowpack, thus creating ice encasement (IE). By decreasing the snowpack insulation capacity and restricting soil-atmosphere gas exchange, modification of the snow properties may lead to colder soil but also to hypoxia and accumulation of trace gases in the subnivean environment. To test the effects of these overwintering conditions changes on plant winter survival and growth, we established a snow manipulation experiment in a coniferous forest in Northern Finland with Norway spruce and Scots pine seedlings. In addition to ambient conditions and prevention of IE, we applied three snow manipulation levels: IE created by artificial rain-on-snow events, snow compaction and complete snow removal. Snow removal led to deeper soil frost during winter, but no clear effect of IE or snow compaction done in early winter was observed on soil temperature. Hypoxia and accumulation of CO2 were highest in the IE plots but, more importantly, the duration of CO2 concentration above 5% was 17 days in IE plots compared to 0 days in ambient plots. IE was the most damaging winter condition for both species, decreasing the proportion of healthy seedlings by 47% for spruce and 76% for pine compared to ambient conditions. Seedlings in all three treatments tended to grow less than seedlings in ambient conditions but only IE had a significant effect on spruce growth. Our results demonstrate a negative impact of winter climate change on boreal forest regeneration and productivity. Changing snow conditions may thus partially mitigate the positive effect of increasing growing season temperatures on boreal forest productivity.

European beech (Fagus sylvatica L.) forests provide multiple essential ecosystem goods and services. The projected climatic conditions for the current century will significantly affect the vitality of European beech. The expected impact of climate change on forest ecosystems will be potentially stronger in southeast Europe than on the rest of the continent. Therefore, our aim was to use the long-term monitoring data of crown vitality indicators in Croatia to identify long-term trends, and to investigate the influence of current and previous year climate conditions and available site factors using defoliation (DEF) and defoliation change (ΔDEF) as response variables. The results reveal an increasing trend of DEF during the study period from 1996 to 2017. In contrast, no significant trend in annual ΔDEF was observed. The applied linear mixed effects models indicate a very strong influence of previous year drought on ΔDEF, while climate conditions have a weak or insignificant effect on DEF. The results suggest that site factors explain 25 to 30% DEF variance, while similar values of conditional and marginal R2 show a uniform influence of drought on ΔDEF. These results suggest that DEF represents the accumulated impact of location-specific stressful environmental conditions on tree vitality, while ΔDEF reflects intense stress and represents the current or recent status of tree vitality that could be more appropriate for analysing the effect of climate conditions on forest trees.

Understanding the relationship between nitrogen (N) availability and stand health in forest ecosystems is crucial, because a large proportion of European forests is subjected to N‐deposition levels beyond their retention capacity. We used data from a long‐term forest monitoring programme (ICP Forests) to test the relationship between an index of N availability, foliar nitrogen : phosphorus (N : P) ratios, tree defoliation and discoloration. We hypothesized a segmented response of stand health to N : P ratios and an improved model‐fit after correcting for climatic covariates. In accordance with the hypothesis, we found a segmented response with a breakpoint for conifer defoliation at N : P ratios as low as 7.3. Inclusion of climatic variables improved the fit of the models, but there was significant collinearity with N : P. Increases in N availability appear, at least for conifers, to have a negative effect on tree health even under N‐limiting conditions. Regulation of N‐deposition levels is consequently as timely as ever. We propose that increases in tree defoliation, other than resulting in serious plant fitness issues, may represent early diagnostic symptoms of N‐addition related imbalances.

Despite being adapted to a wide range of environmental conditions, the vitality of European beech is expected to be significantly affected by the projected effects of climate change, which we attempted to assess with foliar nutrition and crown defoliation, as two different, yet interlinked vitality indicators. Based on 28 beech plots of the ICP Forests Level I network, we set out to investigate the nutritional status of beech in Croatia, the relation of its defoliation and nutrient status, and the effects of environmental factors on this relation. The results indicate a generally satisfactory nutrition of common beech in Croatia. Links between defoliation and nutrition of beech are not very direct or very prominent; differences were observed only in some years and on limited number of plots. However, the applied multinomial logistic regression models show that environmental factors affect the relationship between defoliation and nutrition, as climate and altitude influence the occurrence of differences in foliar nutrition between defoliation categories.

Resource allocation to different plant tissues is likely to be affected by high investment into fruit production during mast years. However, there is a large knowledge gap concerning species-specific differences in resource dynamics. We investigated the influence of mast years on stem growth, leaf production, and leaf carbon (C), nitrogen (N), and phosphorus (P) concentrations and contents in Fagus sylvatica , Quercus petraea , and Q. robur at continental and climate region scales using long-term data from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) and similar datasets. We discussed the results in the light of opposing resource dynamics hypotheses: (i) resource accumulation before mast years and exhaustion after mast years ( resource storage hypothesis ), (ii) shifting resources from vegetative to generative compartments ( resource switching hypothesis ), and (iii) investing resources concurrently in both vegetative and generative compartments ( resource matching hypothesis ). Linear mixed-effects modelling (LMM) showed that both stem growth and leaf production were negatively influenced by weather conditions which simultaneously lead to high fruit production. Thus, the impact of generative on vegetative growth is intermixed with effects of environmental factors. Superposed epoch analyses and LMM showed that for mast behaviour in F. sylvatica , there are indicators supporting the resource storage and the resource switching hypotheses . Before mast years, resources were accumulated, while during mast years resources switched from vegetative to generative tissues with reduced stem and leaf growth. For the Quercus species, stem growth was reduced after mast years, which supports the resource storage hypothesis . LMM showed that leaf C concentrations did not change with increasing fruit production in neither species. Leaf N and P concentrations increased in F. sylvatica , but not in Quercus species. Leaf N and P contents decreased with increasing fruit production in all species, as did leaf C content in F. sylvatica . Overall, our findings suggest different resource dynamics strategies in F. sylvatica and Quercus species, which might lead to differences in their adaptive capacity to a changing climate.