Beech Roots Research Articles

Mode coupling and signal energy distribution in an open-grown European beech tree

The wind-induced response of the branches, the stem, and the roots of an open-grown European beech (Fagus sylvatica L.) was investigated to determine how the airflow’s kinetic energy transferred to the tree is dissipated. The instrumentation in the above-ground tree parts consisted of 69 Tree Response Sensors. The response of the primary roots to the wind load was recorded with six Tree Strain Sensors. The measurements were used to record the tree’s main vibration modes and quantify the energy dissipation from the measurement signals. The signal energy is reduced linearly by 90% from the crown periphery to the stem base. The base of the stem responds to higher wind loads than the branches and the upper part of the stem. The first-order mode frequency band is the narrowest in the roots. To dissipate the kinetic energy transferred to it as efficiently as possible, all tree parts vibrate in overlapping frequency bands in fundamental mode, resulting in mode coupling from the crown periphery to the roots. Mode coupling enables the elastic energy to be transferred as efficiently as possible via the roots to the ground to withstand wind loads and return to the resting position as quickly as possible.

Oriental beech roots improve soil aggregate stability and reduce soil detachment rate in forest lands

In many forestlands, such as Saravan Forest Park, the high capacity of soil detachment from deforestation and subsequent ecological restoration are important issues that need detailed information about plant and tree species for the objectives of soil conservation. To determine if some species are more helpful for the stability of soil aggregates and thus reducing soil erosion, we examined Oriental beech species, compared to three other species (Crataegus ambigue, Primula heterochroma and Rubus persicus) in the Saravan forest park, and measured how root characteristics (root biomass, root length and root weight density) influenced mean weight diameter of aggregates (MWD) and soil detachment rate (Dc). For measuring Dc, undisturbed samples collected from soils under the four studied species and subjected to five slopes (from 4.8 to 31.4%) and five water discharges (from 0.26 to 0.65 l m−1 s−1) by a hydraulic flume. Results showed that Oriental beech roots can significantly influence soil aggregate stability. MWD was greater in soil under Oriental beech species characterized by high values of root biomass, root length and root weight density. Moreover, Dc was significantly lower in soils with Oriental beech species compared to three other species. Pearson's correlation matrix showed a strong positive correlation between MWD and the root characteristics (P < 0.01) (R2 = 0.8). Also, the clustering of soils among the studied species by the principal component analysis (PCA) revealed a clear difference based on the root characteristics and MWD. Generally, this study showed that, compared with other species, the root system of Oriental beech is more effective in the stabilization of slope by the binding effect on the soil, thus improving aggregate stability and reducing soil detachment rates.

Unusual late-fall wildfire in a pre-Alpine Fagus sylvatica forest reduced fine roots in the shallower soil layer and shifted very fine-root growth to deeper soil depth

After an unusual, late-fall wildfire in a European beech (Fagus sylvatica L.) forest in the pre-Alps of northern Italy, the finest roots (0‒0.3 mm diameter) were generally the most responsive to fire, with the effect more pronounced at the shallowest soil depth. While roots 0.3‒1 mm in diameter had their length and biomass at the shallowest soil depth reduced by fire, fire stimulated more length and biomass at the deepest soil depth compared to the control. Fire elevated the total length of dead roots and their biomass immediately and this result persisted through the first spring, after which control and fire-impacted trees had similar fine root turnover. Our results unveiled the fine-root response to fire when subdivided by diameter size and soil depth, adding to the paucity of data concerning fire impacts on beech roots in a natural condition and providing the basis for understanding unusual fire occurrence on root traits. This study suggests that F. sylvatica trees can adapt to wildfire by plastically changing the distribution of fine-root growth, indicating a resilience mechanism to disturbance.

Upside down and the game of C allocation.

Non-structural carbohydrates (NSCs) represent the primary carbon (C) reserves and play a crucial role for plant functioning and resilience. Indeed, these compounds are involved in the regulation between C supply and demand, and in the maintenance of hydraulic efficiency. NSCs are stored in parenchyma of woody organs, which is recognized as a proxy for reserve storage capacity of tree. Notwithstanding the importance of NSCs for tree physiology, their long-term regulation and trade-offs against growth were not deeply investigated. This work evaluated the long-term dynamics of mature tree reserves in stem and root, proxied by parenchyma features, and focusing on the trade off and interplay between the resources allocation in radial growth and reserves in stem and coarse root. In a Mediterranean beech forest, NSCs content, stem and root wood anatomy analysis, and eddy covariance data, were combined. The parenchyma fraction (RAP) of beech root and stem was different, due to differences in axial parenchyma (AP) and narrow ray parenchyma (nRP) fractions. However, these parenchyma components and radial growth showed synchronous inter-annual dynamics between the two organs. In beech stem, positive correlations were found among soluble sugars content and nRP, and among starch content and the AP. Positive correlations were found among Net Ecosystem Exchange (NEE) and AP of both organs. In contrast, NEE was negatively correlated to radial growth of root and stem. Our results suggest a different contribution of stem and roots to reserves storage, and a putative partitioning in the functional roles of parenchyma components. Moreover, a long-term trade-off of C allocation between growth and reserve pool was evidenced. Indeed, in case of C source reduction, trees preferentially allocate C towards reserves pool. Conversely, in high productive years, growth represents the major C sink.

The effects of mixed-species root zones on the resistance of soil bacteria and fungi to long-term experimental and natural reductions in soil moisture

Mixed forest stands tend to be more resistant to drought than species-specific stands partially due to complementarity in root ecology and physiology. We asked whether complementary differences in the drought resistance of soil microbiomes might contribute to this phenomenon. We experimented on the effects of reduced soil moisture on bacterial and fungal community composition in species-specific (single species) and mixed-species root zones of Norway spruce and European beech forests in a 5-year-old throughfall-exclusion experiment and across seasonal (spring-summer-fall) and latitudinal moisture gradients. Bacteria were most responsive to changes in soil moisture, especially members of Rhizobiales, while fungi were largely unaffected, including ectomycorrhizal fungi (EMF). Community resistance was higher in spruce relative to beech root zones, corresponding with the proportions of drought-favored (more in spruce) and drought-sensitive bacterial taxa (more in beech). The spruce soil microbiome also exhibited greater resistance to seasonal changes between spring (wettest) and fall (driest). Mixed-species root zones contained a hybrid of beech- and spruce-associated microbiomes. Several bacterial populations exhibited either enhanced resistance or greater susceptibility to drought in mixed root zones. Overall, patterns in the relative abundances of soil bacteria closely tracked moisture in seasonal and latitudinal precipitation gradients and were more predictive of soil water content than other environmental variables. We conclude that complementary differences in the drought resistance of soil microbiomes can occur and the likeliest form of complementarity in mixed-root zones coincides with the enrichment of drought-tolerant bacteria associated with spruce and the sustenance of EMF by beech.

Implications of hornbeam and beech root systems on slope stability: from field and laboratory measurements to modelling methods

Implications of hornbeam and beech root systems on slope stability: from field and laboratory measurements to modelling methods

Memory Effects of Water Deprivation in European Beech (Fagus sylvatica L.) and Silver Fir (Abies alba Mill.) Seedlings Grown in Mixed Cultivation

Very limited information is available on the drought tolerance of European beech and silver fir in mixed cultivation, both for mature forests and natural regeneration. Particularly, little information is available regarding the significance on memory effects of drought exposure. Therefore, drought memory was analyzed in seedlings of these species grown in mixed cultivation in the present study. The results showed that previous-year drought hardening mediated enhanced biomass accumulation of beech leaves and root in the subsequent year, but did not impact fir growth. Total carbon (C) content was decreased by drought hardening in both the leaves and roots of beech and previous-year needles and roots of fir, in beech probably as a consequence of increased growth. Previous-year drought hardening had no significant effect on relative water contents, total nitrogen (N), or soluble protein contents in leaves and roots of beech and fir, but resulted in decreased total amino acid contents of beech leaves and fir needles. It further reduced structural N in current-year fir needles and decreased C/N ratios in roots of both beech and silver fir seedlings. Generally, the number of interspecific neighbors had no considerable effect on biomass or total C or N contents, as well as N partitioning in leaves and roots of beech and fir seedlings. The present study highlights that drought hardening induces memory effects in European beech and silver fir seedlings in their mixture in the subsequent year of growth, but these memory effects are stronger in beech than in fir.

Land‐use history alters the diversity, community composition and interaction networks of ectomycorrhizal fungi in beech forests

Abstract Forests have expanded across Europe over the last centuries as a consequence of farmland (agricultural and pasture) abandonment. Agricultural practices usually increase soil fertility and reduce the diversity and abundance of ectomycorrhizal (ECM) fungi, essential mutualists of many woody species in temperate and Mediterranean forests. The recovery of this biotic interaction after the cessation of human activities is, thus, crucial for the re‐establishment of functional forest ecosystems. Here we addressed the legacies of past land use and the recovery of the mutualism between ECM fungi and Fagus sylvatica trees in recent beech forests (&lt;50 years) in Northeast Spain. Soil and root samples were collected in six long‐established and eight recent beech forests to analyse soil abiotic properties and the ECM fungi associated with beech roots (Illumina DNA metabarcoding of ECM tips). Up to 609 amplicon sequence variants (ASVs) of ECM fungi were identified, with 220 ASVs found in both forest types. Recent forests had a significantly lower soil organic matter and phosphorus content, which had a significant effect on the community structure of ECM fungi in beech roots. Moreover, beech trees in recent forests interacted with less fungal taxa but had a higher relative abundance of Ascomycota. Tarzetta spp. (Ascomycota, Pezizales) and Lactarius blennius (Basidiomycota, Russulales) emerge as the particular taxa associated with recent and long‐established forests respectively. More specialized mutualistic networks with a lower species normalized degree were found in recent forests, which might result in a lower quality and resilience of the ECM mutualism. Synthesis. Land‐use history modulated the mycorrhizal symbiosis in regenerating beech forests through changes in soil organic matter and nitrogen, which were the main drivers of the differences in fungal community composition and functional types associated with beech trees in recent forests.

Effects of root and leaf litter identity and diversity on oribatid mite abundance, species richness and community composition.

Habitat heterogeneity is an important driver of aboveground species diversity but few studies have investigated effects on soil communities. Trees shape their surrounding by both leaf litter and roots generating small scale heterogeneity and potentially governing community patterns of soil organisms. To assess the role of vegetation for the soil fauna, we studied whether tree species (Fagus sylvatica L., Acer pseudoplatanus L., Fraxinus excelsior L., Tilia cordata Mill.), markedly differing in leaf litter quality and root associated mycorrhizal symbionts, affect oribatid mite communities by shaping below- and aboveground resources and habitat complexity and availability. Oribatid mite abundance, species richness, community structure and the proportion of litter living and parthenogenetic individuals were analyzed and related to microbial biomass and the amount of remaining litter mass. Although leaf litter species with higher nutritional values decomposed considerably faster, microbial biomass only slightly differed between leaf litter species. Neither root species nor leaf litter species affected abundance, species richness or community structure of oribatid mites. However, root species had an effect on the proportion of parthenogenetic individuals with increased proportions in the presence of beech roots. Overall, the results suggest that identity and diversity of vegetation via leaf litter or roots are of minor importance for structuring oribatid mite communities of a temperate forest ecosystem.

Effects of bedrock type and soil chemistry on the fine roots of European beech – A study on the belowground plasticity of trees

We studied the fine root system of Fagus sylvatica in six mature stands on different bedrock down to the rock surface (or to 2 m) to investigate whether (1) the stand total of fine root biomass (FRB) increases, while the fine root live:dead ratio decreases, with decreasing soil base richness, (2) specific root area (SRA) and root tip frequency increase with a decrease in base richness, and (3) FRB is related to profile depth. The three beech stands on deep soil (>2 m profile depth) had on average by 38% larger FRB totals than the stands on shallow soil (60–80 cm), suggesting that limited root space is an important determinant of fine root system size in F. sylvatica. Despite large variation among sites, soil chemistry influenced root morphology only little: fine root diameter depended on soil C/N ratio and root tip frequency on base saturation in a few soil horizons. Much larger morphological differences were found between topsoil and subsoil roots within a profile. We conclude that the fine root system of F. sylvatica varies under similar climatic conditions remarkably little between base-poor and base-rich sites, in contrast to the pronounced topsoil-subsoil differences in root morphology and fine root density.

ATP as Phosphorus and Nitrogen Source for Nutrient Uptake by Fagus sylvatica and Populus x canescens Roots.

The present study elucidated whether roots of temperate forest trees can take up organic phosphorus in the form of ATP. Detached non-mycorrhizal roots of beech (Fagus sylvatica) and gray poplar (Populus x canescens) were exposed under controlled conditions to 33P-ATP and/or 13C/15N labeled ATP in the presence and absence of the acid phosphatase inhibitor MoO42-. Accumulation of the respective label in the roots was used to calculate 33P, 13C and 15N uptake rates in ATP equivalents for comparison reason. The present data shown that a significant part of ATP was cleaved outside the roots before phosphate (Pi) was taken up. Furthermore, nucleotide uptake seems more reasonable after cleavage of at least one Pi unit as ADP, AMP and/or as the nucleoside adenosine. Similar results were obtained when still attached mycorrhizal roots of adult beech trees and their natural regeneration of two forest stands were exposed to ATP in the presence or absence of MoO42-. Cleavage of Pi from ATP by enzymes commonly present in the rhizosphere, such as extracellular acid phosphatases, ecto-apyrase and/or nucleotidases, prior ADP/AMP/adenosine uptake is highly probable but depended on the soil type and the pH of the soil solution. Although uptake of ATP/ADP/AMP cannot be excluded, uptake of the nucleoside adenosine without breakdown into its constituents ribose and adenine is highly evident. Based on the 33P, 13C, and 15N uptake rates calculated as equivalents of ATP the ‘pro and contra’ for the uptake of nucleotides and nucleosides is discussed.Short SummaryRoots take up phosphorus from ATP as Pi after cleavage but might also take up ADP and/or AMP by yet unknown nucleotide transporter(s) because at least the nucleoside adenosine as N source is taken up without cleavage into its constituents ribose and adenine.

Effect of contaminated soil on multitrophic interactions in a terrestrial system

The effect of contaminated soil on multitrophic interactions in a terrestrial system was studied in a mesocosm experiment with European beech, larvae of summer chafer, and entomopathogenic nematodes (EPN). Beech seedlings were grown in non-contaminated forest and garden soil contaminated with potential toxic elements Cd, Pb and Zn, and in the absence/presence of summer chafer larvae. Root morphology and ectomycorrhizal communities were analysed. Volatile organic compounds (VOC) emitted from beech roots were investigated by GC-MS. Humulene, β-caryophyllene, borneol and camphor were selected for chemotaxis assay with EPN Steinernema feltiae, S. carpocapsae and Heterorhabditis bacteriophora. Increased root tip density was observed as a response to soil contamination and root herbivory. Soil contamination decreased ectomycorrhizal species richness. Analyses of VOC yielded 14 different compounds. Results showed that the least mobile nematode species towards the VOC tested was S. feltiae. β-caryophyllene was an attractant for S. carpocapsae and humulene a weak attractant for H. bacteriophora. Beech roots emitted VOC that affected movement of EPN. β-caryophyllene was detected in beech roots regardless of treatment, indicating that VOC that affect movement of EPN are emitted even in the absence of direct root herbivore attack.

Nitrogen Nutrition of European Beech Is Maintained at Sufficient Water Supply in Mixed Beech-Fir Stands

Research highlights: Interaction effects of coniferous on deciduous species have been investigated before the background of climate change. Background and objectives: The cultivation of European beech (Fagus sylvatica L.) in mixed stands has currently received attention, since the future performance of beech in mid-European forest monocultures in a changing climate is under debate. We investigated water relations and nitrogen (N) nutrition of beech in monocultures and mixed with silver-fir (Abies alba Mill.) in the Black Forest at different environmental conditions, and in the Croatian Velebit at the southern distribution limit of beech, over a seasonal course at sufficient water availability. Material and methods: Water relations were analyzed via δ13C signatures, as integrative measures of water supply assuming that photosynthesis processes were not impaired. N nutrition was characterized by N partitioning between soluble N fractions and structural N. Results: In the relatively wet year 2016, water relations of beech leaves, fir needles and roots differed by season, but generally not between beech monocultures and mixed cultivation. At all sites, previous and current year fir needles revealed significantly lower total N contents over the entire season than beech leaves. Fir fine roots exhibited higher or similar amounts of total N compared to needles. Correlation analysis revealed a strong relationship of leaf and root δ13C signatures with soil parameters at the mixed beech stands, but not at pure beech stands. While glutamine (Gln) uptake capacity of beech roots was strongly related to soil N in the monoculture beech stands, arginine (Arg) uptake capacities of beech roots were strongly related to soil N in mixed stands. Conclusions: Leaf N contents indicated a facilitative effect of silver-fir on beech on sites where soil total N concentrations where low, but an indication of competition effect where it was high. This improvement could be partially attributed to protein contents, but not to differences in uptake capacity of an individual N source. From these results it is concluded that despite similar performance of beech trees at the three field sites investigated, the association with silver-fir mediated interactive effects between species association, climate and soil parameters even at sufficient water supply.

Early-stage of invasion by beech bark disease does not necessarily trigger American beech root sucker establishment in hardwood stands

Two concomitant phenomena currently affect the dynamics of sugar maple-American beech (AB) stands in northeastern North America: beech bark disease (BBD), and increased AB understory density. Many studies suggest a causal link between the two phenomena, i.e., BBD favouring beech regeneration. But this link has yet to be experimentally demonstrated. To address the question, we compared regeneration composition between recently BBD-affected and -unaffected stands. A total of 109 stands were sampled; half were affected by BBD. Seedling and sapling density were assessed, together with the origin (seedling or sprout). While BBD affects stands in the eastern part of the study region, AB was observed in the understory across the entire region. No clear difference in AB sprout density between BBD-affected and -unaffected stands was observed while AB seedling density—as well as pooled AB seedling and sprout density were higher in unaffected stands. Findings suggests that BBD, in its early stage, is not a necessary trigger of AB understory establishment. Yet, AB sapling basal area generally was higher in stands affected by BBD, likely indicating a greater rate of AB understory development due to increased light availability beneath a more open crown canopy. That development can lead to AB understory dominance. This distinction—BBD not necessarily triggering AB root sucker establishment but favoring AB advance regeneration development—also questions the generalized perception that dense AB thickets necessarily originate from root suckers.

Affiliation of Dihydrolipoyl Dehydrogenase Allozymes in Mycorrhizae of European Forest Trees and Characterization of the Enzyme of the Matt Bolete (&amp;lt;i&amp;gt;Xerocomus pruinatus&amp;lt;/i&amp;gt;) and the Bay Bolete (&amp;lt;i&amp;gt;X. badius&amp;lt;/i&amp;gt;)

Mycorrhizal roots of the deciduous trees European beech (Fagus sylvatica (L.)) and Sessile oak (Quercus petraea (MattuschkaLiebl.)) and the conifers Norway spruce (Picea abies (L.) H. Karst.) and European larch (Larix decidua (Mill.)) associated with the ectomycorrhizal fungi matt bolete (Xerocomus pruinatus (Fries 1835)) or bay bolete (X. badius (Fries 1818)) were analysed with respect to the occurrence of dihydrolipoyl dehydrogenase (EC 1.8.1.4) allozymes. In root tissues of the two deciduous trees, two gene loci could be visualized after cellulose acetate electrophoresis while three loci were expressed in root tissues of the two coniferous species. The two fungal species and further ectomycorrhizal fungi expressed exclusively one dihydrolipoyl dehydrogenase gene. In Xerocomus pruinatus and X. badius, the dihydrolipoyl dehydrogenase gene consists of 1460 bp and 1370 bp, respectively, including five introns each consisting of 52 bp. Their DNA sequences correspond to 70 to 90% to other fungal dihydrolipoyl dehydrogenase genes. One monomer of the dimeric dihydrolipoyl dehydrogenase enzyme consists of 486 (X. pruinatus) or 454 (X. badius) amino acids which sum up to a molecular mass of 55 kDa (X. pruinatus), respectively 52 kDa (X. badius). The number of positively charged amino acid residues makes 79 (X. pruinatus) and 68 (X. badius) and the number of negatively charged amino acid residues was calculated to make 46 (X. pruinatus) and 48 (X. badius); isoelectric points make 9.99 (X. pruinatus) and 9.68 (X. badius). Calculated three dimensional structures reveal a short NADH binding site being part of a larger FAD-binding site and a binding/dimerization domain.

Beech shearing/high-stumping to control understory interference

The research evaluated opportunities for cutting off small understory beech root suckers at 0.6-0.9 m above ground. By six years (pilot test) after cutting with lopping shears in a closed-canopy uneven-aged northern hardwood stand, 91% of high stumps lacking a residual live branch had died. Of those cut above the lowest living branch, all survived. In a later operational trial by contract crews using chainsaws two growing seasons before a selection system cutting, 75% of high stumps without a residual live branch had died by the fifth year. Of stumps cut below the lowest living branch and that initially sprouted, only 7% had a live sprout by the sixth year. Among high stumps cut above the lowest branch, 84% survived.

Immunodepletion of α‐plurivorin effector leads to loss of virulence of Phytophthora plurivora towards Fagus sylvatica

SummaryPhytophthora species secrete several proteins during their interaction with plants. Some of these proteins manipulate host metabolism favouring infection, while others can be recognized by plants thus triggering defence. Elicitins are known to elicit plant defences, leading to resistance. Here, we characterized the elicitin α‐plurivorin and proved that it was essential for the virulence of Phytophthora plurivora towards Fagus sylvatica. The immunodepletion of this peptide impaired its penetration into host tissue and in parallel P. plurivora lost its ability to colonize beech roots. Furthermore, the lack of α‐plurivorin inside the host led to an up‐regulation of several defence‐related genes of both salicylic acid and jasmonate/ethylene pathways, suggesting that α‐plurivorin might act as an effector‐triggered susceptibility during infection. Consequently, plants survived infection with P. plurivora after α‐plurivorin immunodepletion, whereas the majority of the infected control plants had died at the end of the experiment. Because canonical elicitins are ubiquitously secreted by many Phytophthora species, it is possible that these molecules may play a similar role in other susceptible interactions, being a potential target for controlling Phytophthora diseases.

Influence of Root Diameter and Soil Depth on the Xylem Anatomy of Fine- to Medium-Sized Roots of Mature Beech Trees in the Top- and Subsoil.

Despite their importance for water uptake and transport, the xylem anatomical and hydraulic properties of tree roots have only rarely been studied in the field. We measured mean vessel diameter (D), vessel density (VD), relative vessel lumen area (lumen area per xylem area) and derived potential hydraulic conductivity (Kp) in the xylem of 197 fine- to medium-diameter roots (1–10 mm) in the topsoil and subsoil (0–200 cm) of a mature European beech forest on sandy soil for examining the influence of root diameter and soil depth on xylem anatomical and derived hydraulic traits. All anatomical and functional traits showed strong dependence on root diameter and thus root age but no significant relation to soil depth. Averaged over topsoil and deep soil and variable flow path lengths in the roots, D increased linearly with root diameter from ∼50 μm in the smallest diameter class (1–2 mm) to ∼70 μm in 6–7 mm roots (corresponding to a mean root age of ∼12 years), but remained invariant in roots >7 mm. D never exceeded ∼82 μm in the 1–10 mm roots, probably in order to control the risk of frost- or drought-induced cavitation. This pattern was overlain by a high variability in xylem anatomy among similar-sized roots with Kp showing a higher variance component within than between root diameter classes. With 8% of the roots exceeding average Kp in their diameter class by 50–700%, we obtained evidence of the existence of ‘high-conductivity roots’ indicating functional differentiation among similar-sized roots. We conclude that the hydraulic properties of small to medium diameter roots of beech are mainly determined by root age, rendering root diameter a suitable predictor of hydraulic functioning, while soil depth – without referring to path length – had a negligible effect.

Changes in the fine root proteome of Fagus sylvatica L. trees associated with P-deficiency and amelioration of P-deficiency

Phosphorus is often the least available macronutrient in soil. Lack in phosphorus has detrimental effect on growth and biomass production of European Fagus sylvatica L., a major trees species in temperate forests. In contrast to leaf tissues, few studies have examined changes in the root system and no study has ever investigated the proteomic changes affected in beech roots by a lack in available phosphate (P). Here, we studied roots of young Fagus sylvatica L. trees in their native soils from two forests sites with contrasting availability of P: one P rich and P poor soil. To understand also the response to P fertilization, the trees were fertilized with triple superphosphate and the proteome of fine roots of all conditions was compared. Gel-free mass-spectrometry-based shotgun proteomics revealed that the proteome was differentially affected by diverging P availabilities. The proteomic changes that took place as the result of P fertilization were dependent on the supply level of P before the fertilization. When P was supplied to the P-rich soil proteins related to cell biogenesis exhibited increased abundances. Addition of P to soil that was strongly limited in P resulted in increased abundance of proteins associated with amino acid metabolism and transport. Biological significanceBeech (Fagus sylvatica L.) forests have a huge ecological and economic value across Europe. In recent years, however, these forest sites increasingly suffer under phosphorus (P) deficiency. As the consequence, growth and vitality of beech forests is impaired. For this reason, this study was conducted with the aim to identify and understand proteomic impairments and adjustments that evolve in the fine roots under both, a P deficiency and an amelioration thereof. For this, we analyzed (1) the fine root proteome of young beech trees grown (2) at two soil sites that contrast in their degree of availability P (low vs. high) in dependency (3) to a fertilization with P. This experiment revealed fundamental differences with respect to proteomic changes in dependency on the severity of P limitation and helped to identify processes that take place after amelioration of the deficiency. This information is useful to understand which physiological processes are impaired under P deficiency and, thus, impair growth. The fertilization experiment enabled to identify developmental processes that take place in fine roots when concentration of available P was increased. They are “cellular component organization and biogenesis” in the P rich soil and “synthesis of organonitrogen-containing compounds” in the P poor soil.

The Deep Root System of Fagus sylvatica on Sandy Soil: Structure and Variation Across a Precipitation Gradient

When applied to climate change-related precipitation decline, the optimal partitioning theory (OPT) predicts that plants will allocate a larger portion of carbon to root growth to enhance the capacity to access and acquire water. However, tests of OPT applied to the root system of mature trees or stands exposed to long-term drying show mixed, partly contradicting, results, indicating an overly simplistic understanding of how moisture affects plant-internal carbon allocation. We investigated the response of the root system (0–240 cm depth) of European beech to long-term decrease in water supply in six mature forests located across a precipitation gradient (855–576 mm mean annual precipitation, MAP). With reference to OPT, we hypothesized that declining precipitation across this gradient would: (H1) cause the profile total of fine root biomass (FRB; roots <2 mm) to increase relative to total leaf mass; (H2) trigger a shift to a shallower root system; and (H3) induce different responses in the depth distributions of different root diameter classes. In contradiction to H1, neither total FRB (0–240 cm) nor the FRB:leaf mass ratio changed significantly with the MAP decrease. The support for H2 was only weak: the 95% rooting depth of fine roots decreased with decreasing MAP, whereas the maximum extension of small coarse roots (2–5 mm) increased, indicating contrasting responses of different root diameter classes. We conclude that long-term decline in water supply leads to only minor adaptive modification with respect to the size and structure of the beech root system, with notable change in the depth extension of some root diameter classes but limited capacity to alter the fine root:leaf mass ratio. It appears that OPT cannot adequately predict C allocation shifts in mature trees when exposed to long-term drying.