Effects of air humidity and soil moisture on secondary metabolites in the leaves and roots of Betula pendula of different competitive status.

Plant secondary metabolites (PSMs) defend plants against abiotic stresses, including those caused by climate change and against biotic stresses, such as herbivory and competition. There is a trade-off between allocating available carbon to growth and defence in stressful environments. However, our knowledge about trade-off is limited, especially when abiotic and biotic stresses co-occur. We aimed to understand the combined effect of increasing precipitation and humidity, the tree's competitive status, and canopy position on leaf secondary metabolites (LSMs) and fine root secondary metabolites (RSMs) in Betula pendula. We sampled 8-year-old B. pendula trees growing in the free air humidity manipulation (FAHM) experimental site, where treatments included elevated relative air humidity and elevated soil moisture. A high-performance liquid chromatography-quadrupole-time of flight mass spectrometer (HPLC-qTOF-MS) was used to analyse secondary metabolites. Our results showed accumulation of LSM depends on the canopy position and competitive status. Flavonoids (FLA), dihydroxybenzoic acids (HBA), jasmonates (JA) and terpene glucosides (TG) were higher in the upper canopy, and FLA, monoaryl compounds (MAR) and sesquiterpenoids (ST) were higher in dominant trees. The FAHM treatments had a more distinct effect on RSM than on LSM. The RSMs were lower in elevated air humidity and soil moisture conditions than in control conditions. The RSM content depended on the competitive status and was higher in suppressed trees. Our study suggests that young B. pendula will allocate similar amounts of carbon to constitutive chemical leaf defence, but a lower amount to root defence (per fine root biomass) under higher humidity.

Climate and Competitive Status Modulate the Variation in Secondary Metabolites More in Leaves Than in Fine Roots of Betula pendula.

Plant secondary metabolites have many important functions; they also determine the productivity and resilience of trees under climate change. The effects of environmental factors on secondary metabolites are much better understood in above-ground than in below-ground part of the tree. Competition is a crucial biotic stress factor, but little is known about the interaction effect of climate and competition on the secondary chemistry of trees. Moreover, competition effect is usually overlooked when analyzing the sources of variation in the secondary chemistry. Our aim was to clarify the effects of competitive status, within-crown light environment, and climate on the secondary chemistry of silver birch (Betula pendula Roth). We sampled leaves (from upper and lower crown) and fine roots from competitively dominant and suppressed B. pendula trees in plantations along a latitudinal gradient (56-67° N) in Fennoscandia, with mean annual temperature (MAT) range: -1 to 8°C. Secondary metabolites in leaves (SML) and fine roots (SMFR) were determined with an HPLC-qTOF mass spectrometer. We found that SML content increased significantly with MAT. The effect of competitive stress on SML strengthened in colder climates (MAT<4°C). Competition and shade initiated a few similar responses in SML. SMFR varied less with MAT. Suppressed trees allocated relatively more resources to SML in warmer climates and to SMFR in colder ones. Our study revealed that the content and profile of secondary metabolites (mostly phenolic defense compounds and growth regulators) in leaves of B. pendula varied with climate and reflected the trees' defense requirements against herbivory, exposure to irradiance, and competitive status (resource supply). The metabolic profile of fine roots reflected, besides defense requirements, also different below-ground competition strategies in warmer and colder climates. An increase in carbon assimilation to secondary compounds can be expected at northern latitudes due to climate change.

The initial overreaction of carbon cycle to elevated atmospheric humidity levels off over time - a FAHM study in a young birch forest.

Ecosystem responses to climate change are mainly predicted based on short-term studies. However, the first response can be a temporary overreaction, different from the later response of the more acclimated ecosystem. The current paper is a follow-up study of our previous article, where the effect of elevated atmospheric humidity on forest ecosystem carbon (C) balance was studied in a young silver birch (Betula pendula Roth) forest after two years of humidification. Here, we present the C balance of the same forest measured two years later when humidification treatment had been performed for four years. We revealed that the higher C sequestration capacity of the humidified birch forest ecosystem was an initial overreaction, which levelled off after four years of humidification, when the ecosystem became more acclimated to wetter conditions. Understorey production reacted rapidly and strongly by increasing belowground production more than twofold, but this reaction ceased after four years of humidification treatment. Trees responded to a lesser extent, and the initially decreased aboveground growth was recovered after four years of humidification, when the biomass allocation to tree fine-roots was increased. Our results showed that at early forest age, understorey plant production dominated in the whole ecosystem C sequestration capacity. But in the later stage, the most important C sink was biomass production of birches, and since the tree biomass production no longer differed between the treatments, C sequestration of the whole ecosystem did not differ either. The findings confirm that a preliminary reaction of an ecosystem can be different from the later response, which needs to be taken into account when prognosing the climate change consequences for carbon sequestration.

Adaptive root foraging strategies along a boreal-temperate forest gradient.

The tree root-mycorhizosphere plays a key role in resource uptake, but also in the adaptation of forests to changing environments. The adaptive foraging mechanisms of ectomycorrhizal (EcM) and fine roots of Picea abies, Pinus sylvestris and Betula pendula were evaluated along a gradient from temperate to subarctic boreal forest (38 sites between latitudes 48°N and 69°N) in Europe. Variables describing tree resource uptake structures and processes (absorptive fine root biomass and morphology, nitrogen (N) concentration in absorptive roots, extramatrical mycelium (EMM) biomass, community structure of root-associated EcM fungi, soil and rhizosphere bacteria) were used to analyse relationships between root system functional traits and climate, soil and stand characteristics. Absorptive fine root biomass per stand basal area increased significantly from temperate to boreal forests, coinciding with longer and thinner root tips with higher tissue density, smaller EMM biomass per root length and a shift in soil microbial community structure. The soil carbon (C) : N ratio was found to explain most of the variability in absorptive fine root and EMM biomass, root tissue density, N concentration and rhizosphere bacterial community structure. We suggest a concept of absorptive fine root foraging strategies involving both qualitative and quantitative changes in the root-mycorrhiza-bacteria continuum along climate and soil C : N gradients.

Elevated Air Humidity Changes Soil Bacterial Community Structure in the Silver Birch Stand.

Soil microbes play a fundamental role in forest ecosystems and respond rapidly to changes in the environment. Simultaneously with the temperature increase the climate change scenarios also predict an intensified hydrological cycle for the Baltic Sea runoff region. The aim of this study was to assess the effect of elevated air humidity on the top soil microbial community structure of a silver birch (Betula pendula Roth.) stand by using a free air humidity manipulation facility (FAHM). The bacterial community structures of bulk soil and birch rhizosphere were analyzed using high-throughput sequencing of bacteria-specific16S rRNA gene fragments and quantification of denitrification related genes. The increased air humidity altered both bulk soil and rhizosphere bacterial community structures, and changes in the bacterial communities initiated by elevated air humidity were related to modified soil abiotic and biotic variables. Network analysis revealed that variation in soil bacterial community structural units is explained by altered abiotic conditions such as increased pH value in bulk soil, while in rhizosphere the change in absorptive root morphology had a higher effect. Among root morphological traits, the absorptive root diameter was strongest related to the bacterial community structure. The changes in bacterial community structures under elevated air humidity are associated with shifts in C, N, and P turnover as well as mineral weathering processes in soil. Increased air humidity decreased the nir and nosZ gene abundance in the rhizosphere bacterial community. The potential contribution of the denitrification to the N2O emission was not affected by the elevated air humidity in birch stand soil. In addition, the study revealed a strong link between the bacterial community structure, abundance of denitrification related genes, and birch absorptive root morphology in the ecosystem system adaptation to elevated air humidity.

Elevated air humidity affects hydraulic traits and tree size but not biomass allocation in young silver birches (Betula pendula).

As changes in air temperature, precipitation, and air humidity are expected in the coming decades, studies on the impact of these environmental shifts on plant growth and functioning are of major importance. Greatly understudied aspects of climate change include consequences of increasing air humidity on forest ecosystems, predicted for high latitudes. The main objective of this study was to find a link between hydraulic acclimation and shifts in trees' resource allocation in silver birch (Betula pendula Roth) in response to elevated air relative humidity (RH). A second question was whether the changes in hydraulic architecture depend on tree size. Two years of application of increased RH decreased the biomass accumulation in birch saplings, but the biomass partitioning among aboveground parts (leaves, branches, and stems) remained unaffected. Increased stem Huber values (xylem cross-sectional area to leaf area ratio) observed in trees under elevated RH did not entail changes in the ratio of non-photosynthetic to photosynthetic tissues. The reduction of stem-wood density is attributable to diminished mechanical load imposed on the stem, since humidified trees had relatively shorter crowns. Growing under higher RH caused hydraulic conductance of the root system (K R) to increase, while K R (expressed per unit leaf area) decreased and leaf hydraulic conductance increased with tree size. Saplings of silver birch acclimate to increasing air humidity by adjusting plant morphology (live crown length, slenderness, specific leaf area, and fine-root traits) and wood density rather than biomass distribution among aboveground organs. The treatment had a significant effect on several hydraulic properties of the trees, while the shifts were largely associated with changes in tree size but not in biomass allocation.

Morphological plasticity of ectomycorrhizal short roots in Betula sp and Picea abies forests across climate and forest succession gradients: its role in changing environments.

Morphological plasticity of ectomycorrhizal (EcM) short roots (known also as first and second order roots with primary development) allows trees to adjust their water and nutrient uptake to local environmental conditions. The morphological traits (MTs) of short-living EcM roots, such as specific root length (SRL) and area, root tip frequency per mass unit (RTF), root tissue density, as well as mean diameter, length, and mass of the root tips, are good indicators of acclimation. We investigated the role of EcM root morphological plasticity across the climate gradient (48-68°N) in Norway spruce (Picea abies (L.) Karst) and (53-66°N) birch (Betula pendula Roth., B. pubescens Ehrh.) forests, as well as in primary and secondary successional birch forests assuming higher plasticity of a respective root trait to reflect higher relevance of that characteristic in acclimation process. We hypothesized that although the morphological plasticity of EcM roots is subject to the abiotic and biotic environmental conditions in the changing climate; the tools to achieve the appropriate morphological acclimation are tree species-specific. Long-term (1994-2010) measurements of EcM roots morphology strongly imply that tree species have different acclimation-indicative root traits in response to changing environments. Birch EcM roots acclimated along latitude by changing mostly SRL [plasticity index (PI) = 0.60], while spruce EcM roots became adjusted by modifying RTF (PI = 0.68). Silver birch as a pioneer species must have a broader tolerance to environmental conditions across various environments; however, the mean PI of all MTs did not differ between early-successional birch and late-successional spruce. The differences between species in SRL, and RTF, diameter, and length decreased southward, toward temperate forests with more favorable growth conditions. EcM root traits reflected root-rhizosphere succession across forest succession stages.