Insolation and disturbance history drive biocrust biodiversity in Western Montana rangelands

Background and Aims

Biological soil crust (biocrust) communities, though common and important in the intermountain west, have received little research attention. There are gaps in understanding what influences biocrust species’ abundance and distributions in this ecoregion. Climatic, edaphic, topographic, and biotic forces, in addition to anthropogenic disturbance can all influence the biocrust.

Methods

We determined the relative influence of several possible environmental filters in biocrust communities of western Montana (USA) grasslands at two spatial scales. The larger scale exploited strong topographically-dictated climatic variation across >60km², while the smaller scale focused on differences among distinct microsites within ~700m² plots.

Results

We detected a total of 96 biocrust taxa, mostly lichens. Biocrust richness at each site ranged from 0 to 39 species, averaging 14 species. Insolation, aspect, and disturbance history were the strongest predictors of biocrust richness, abundance, and species turnover across the landscape; soil texture was influential for some biocrust community properties. Steep, north-facing slopes that receive longer periods of shade harbored higher diversity and cover of biocrust than south-facing sites. At a small scale, interspaces among native herbaceous communities supported the greatest diversity of biocrust species, but microsites under shrub canopies supported the greatest cover.

Conclusions

We found that, among the variables investigated, tillage, insolation, soil texture and the associated vegetation community were the most important drivers of biocrust abundance and species richness. This study can inform the practice of restoration and conservation, and also guide future work to improve predictions of biocrust properties.

     

Biocrusts enhance soil fertility and <Emphasis Type="Italic">Bromus tectorum</Emphasis> growth,and interact with warming to influence germination

Background and aims

Biocrusts are communities of cyanobacteria, mosses, and/or lichens found in drylands worldwide. Biocrusts are proposed to enhance soil fertility and productivity, but simultaneously act as a barrier to the invasive grass, Bromus tectorum, in western North America. Both biocrusts and B. tectorum are sensitive to climate change drivers, yet how their responses might interact to affect dryland ecosystems is unclear.

Methods

Using mesocosms with bare soil versus biocrust cover, we germinated B. tectorum seeds collected from warmed, warmed + watered, and ambient temperature plots within a long-term climate change experiment on the Colorado Plateau, USA. We characterized biocrust influences on soil fertility and grass germination, morphology, and chemistry.

Results

Biocrusts increased soil fertility and B. tectorum biomass, specific leaf area (SLA), and root:shoot ratios. Germination rates were unaffected by mesocosm cover-type. Biocrusts delayed germination timing while also interacting with the warmed treatment to advance, and with the warmed + watered treatment to delay germination.

Conclusions

Biocrusts promoted B. tectorum growth, likely through positive influence on soil fertility which was elevated in biocrust mesocosms, and interacted with seed treatment-provenance to affect germination. Understanding how anticipated losses of biocrusts will affect invasion dynamics will require further investigation of how plant plasticity/adaptation to specific climate drivers interact with soil and biocrust properties.

     

Trade-off between C and N recycling and N<Subscript>2</Subscript>O emissions of soils with summer cover crops in subtropical agrosystems

Aims

Biological soil crusts (biocrusts) are soil-surface communities in drylands, dominated by cyanobacteria, mosses, and lichens. They provide key ecosystem functions by increasing soil stability and influencing soil hydrologic, nutrient, and carbon cycles. Because of this, methods to reestablish biocrusts in damaged drylands are needed. Here we test the reintroduction of field-collected vs. greenhouse-cultured biocrusts for rehabilitation.

Methods

We collected biocrusts for 1) direct reapplication, and 2) artificial cultivation under varying hydration regimes. We added field-collected and cultivated biocrusts (with and without hardening treatments) to bare field plots and monitored establishment.

Results

Both field-collected and cultivated cyanobacteria increased cover dramatically during the experimental period. Cultivated biocrusts established more rapidly than field-collected biocrusts, attaining ~82% cover in only one year, but addition of field-collected biocrusts led to higher species richness, biomass (as assessed by chlorophyll a) and level of development. Mosses and lichens did not establish well in either case, but late successional cover was affected by hardening and culture conditions.

Conclusions

This study provides further evidence that it is possible to culture biocrust components from later successional materials and reestablish cultured organisms in the field. However, more research is needed into effective reclamation techniques.

     

Crop mixtures: does niche complementarity hold for belowground resources? An experimental test using rice genotypic pairs

Aims

A growing body of research supports the feasibility of biocrust rehabilitation. Identifying populations of key species that are amenable to cultivation and that are resilient in rehabilitation contexts would advance the efficacy of these technologies. Here we investigate the growth and stress response of the cosmopolitan biocrust moss, Syntrichia ruralis.

Methods

We sampled populations of S. ruralis along a precipitation seasonality gradient from the Colorado Plateau ecoregion of the western United States. We cultivated these populations in an experiment manipulating duration of hydration periods on a weekly cycle. We then treated greenhouse grown materials with brief, stressful watering events, measuring how many events they could survive.

Results

All populations grew at an accelerated rate compared to growth in a natural setting, at least doubling biomass in five months. Increasing duration of hydration periods led to more growth in all but one population. Volunteer biocrust algae and cyanobacteria developed during cultivation, and differed among populations. Greenhouse grown mosses differed in their response to stressful watering, with the most susceptible populations dying at half the number events compared to the most tolerant.

Conclusions

These findings argue for informed selection and deployment of Syntrichia ruralis populations for soil rehabilitation.

     

Carbon mineralization and microbial activity in agricultural topsoil and subsoil as regulated by root nitrogen and recalcitrant carbon concentrations

Background and aims

Biocrust morphology is often used to infer ecological function, but morphologies vary widely in pigmentation and thickness. Little is known about the links between biocrust morphology and the composition of constituent microbial community. This study aimed to examine these links using dryland crusts varying in stage and morphology.

Methods

We compared the microbial composition of three biocrust developmental stages (Early, Mid, Late) with bare soil (Bare) using high Miseq Illumina sequencing. We used standard diversity measures and network analysis to explore how microbe-microbe associations changed with biocrust stage.

Results

Biocrust richness and diversity increased with increasing stage, and there were marked differences in the microbial signatures among stages. Bare and Late stages were dominated by Alphaproteobacteria, but Cyanobacteria was the dominant phylum in Early and Mid stages. The greatest differences in microbial taxa were between Bare and Late stages. Network analysis indicated highly-connected hubs indicative of small networks.

Conclusions

Our results indicate that readily discernible biocrust features may be good indicators of microbial composition and structure. These findings are important for land managers seeking to use biocrusts as indicators of ecosystem health and function. Treating biocrusts as a single unit without considering crust stage is likely to provide misleading information on their functional roles.

     

Threshold effect of the groundwater depth on the photosynthetic efficiency of <Emphasis Type="Italic">Tamarix chinensis</Emphasis> in the Yellow River Delta

Background and aims

Functional traits may underlie differences in niches, which promote plant species co-existence, but also differences in competitive ability, which drive competitive exclusion. Empirical evidence concerning the contribution of different traits to niche differentiation and the ability to supress and tolerate competitors is very limited, particularly when considering belowground interactions.

Methods

We grew 26 temperate grassland species along a density gradient of interspecific competitors to determine which belowground traits a) explain species’ ability to suppress and tolerate neighbours and b) contribute to niche differentiation, such that species with dissimilar trait values experience reduced competition.

Results

We found that having larger root systems with extensive horizontal spread and lower root tissue density enabled efficient suppression of neighbours but did not significantly contribute to the ability to tolerate competition. Species with deeper root systems, lower specific root length and less branched roots were better at tolerating competition, but these traits did not significantly affect the ability to suppress neighbours. None of the measured traits contributed significantly to niche differentiation, either individually or in combination.

Conclusions

This study provides little support for belowground traits contributing to species co-existence through niche differentiation. Instead, different sets of weakly correlated traits enable plants to either suppress or tolerate their competitors.

     

Root morphology acclimation to phosphorus supply by six cultivars of <Emphasis Type="Italic">Trifolium subterraneum</Emphasis> L

Aims

Trifolium subterraneum L. is the predominant annual pasture legume in southern Australia. Cultivars with improved phosphorus (P) foraging ability would improve the P-use efficiency of agricultural systems. We therefore investigated variation in root traits related to P-uptake among six cultivars.

Methods

Micro-swards were grown at six levels of P in field soil with indigenous arbuscular mycorrhizal (AM) fungi for six weeks. Dry matter yield, tissue P concentration, rhizosphere carboxylates, AM fungal colonisation and root morphological traits were measured.

Results

The cultivars showed similar shoot and root yield responses to P supply. Average root diameter did not change, specific root length (SRL) increased and root tissue density (RTD) decreased with increased P supply. Amounts of total rhizosphere carboxylates were low (<1.2 nmol cm^?1 root). The percentage of root length colonised by AM fungi was greatest (29–43 %) at an intermediate level (8 mg kg^?1 dry soil) of P supply.

Conclusions

Most differences among cultivars were reasonably consistent across P supply levels, indicating greater numbers of lines could be screened reliably at a single P level. Low colonisation by AM fungi at low P supply deserves consideration when selecting soils for cultivar comparisons. Increased SRL and decreased RTD at high P supply likely result from self-shading within the micro-swards and warrant further investigation.

     

Establishing rates of lateral expansion of cyanobacterial biological soil crusts for optimal restoration

Aims

Biocrusts that form on topsoils contribute ecosystem services to drylands, and their loss under anthropogenic pressure has negative ecological consequences. Therefore, development of biocrust inoculation technology for restoration is of interest. This requires knowledge of biocrust growth and dispersal. To contribute to this, we determined the speed at which biocrusts expand laterally based on the self-propelled motility of cyanobacteria.

Methodology

We inoculated sterile soil with natural biocrusts and incubated them over a year in a greenhouse under conditions mimicking local precipitation, monitoring the crust’s lateral expansion using time-course photography, chlorophyll a content, and microscopic inspection. Concurrent uninoculated controls served to monitor, and discount, natural inoculation by aeolian propagules.

Results

While the expansion front was highly variable in space, biocrusts expanded in the order of 2 cm month^?1, but only in seasons with moderate temperatures (Spring and Fall). Microcoleus vaginatus, Microcoleus steenstrupii, and Scytonema spp. advanced at averages of 1 cm month^?1, the crust advance front being preferentially driven by specialized propagules (hormogonia). These rates are within expectations based on instantaneous gliding motility speeds of cyanobacteria.

Conclusions

Based on the expansion capability of biocrusts during growth seasons, greenhouse inoculum units can be optimally spaced to fill 4–8 cm gaps.

     

Dynamic succession of chickpea rhizobia over years and sampling sites in Xinjiang,China

Aims

Cyanobacterial and algal communities have a large effect on biocrust formation and development. Biocrust species and abundance vary spatially and temporally due to different environmental factors. The relationships among cyanobacterial and algal communities, and biocrust function have been studied extensively. Lacking, however, are studies of temporal changes in a similar landscape where environmental conditions are similar, but where biocrust formation is different.

Methods

Biocrusts of different ages were located in the Loess Plateau in an area that had experienced a landslide. We examined changes in cyanobacterial and algal communities, carbon, nutrients, and the composition of dissolved organic matter in the topsoil, and the relationships among the community and soil characteristics using redundancy analysis.

Results

Phormidium tenue (Cyanophyta) dominated in all biocrusts, and co-dominated in a newly formed crust with Euglena sp. (Euglenophyta). Oscillatoria sp. (Cyanophyta) increased with biocrust age. Oscillatoria was positively correlated with carbon fixation and nutrient (nitrogen and phosphorus) accumulation in topsoils.

Conclusions

While incubation of Phormidium tenue and Euglena sp. is suggested for rapid biocrust formation at an early stage, increasing the abundance of Phormidium tenue and Oscillatoria sp. can promote carbon fixation and nutrient (nitrogen and phosphorus) accumulation, thereby accelerating biocrusts into a later stage.

     

Imbalanced plant stoichiometry at contrasting geologic-derived phosphorus sites in subtropics: the role of microelements and plant functional group

Aims

In western North America ectomycorrhizal fungi are critical to establishment of conifers in low nitrogen soils. Fire can affect both ectomycorrhizal fungi and soil properties, and inoculation with ectomycorrhizal fungi is recommended when planting on burns for restoration. The aim of this study was to examine how Suillus species used in inoculation affect whitebark pine (Pinus albicaulis L.) seedlings planted in fire-impacted soil.

Methods

In a greenhouse experiment, Suillus-colonized and uncolonized whitebark pine seedlings were planted in unsterilized and sterilized (control) soil from a recent burn. After 6 months, foliar nitrogen and carbon content, concentration, and stable isotope values were assessed, along with growth parameters.

Results

When seedlings were colonized, biomass was 61% greater, foliar nitrogen content 25% higher, foliar nitrogen concentration 30–63% lower; needles had lower δ¹⁵N and higher δ¹³C. Differences were more pronounced in sterilized soil where colonization was higher. Foliar N content was negatively correlated with δ¹⁵N values.

Conclusions

Colonization by host-specific fungi produced larger seedlings with higher foliar nitrogen content in both burn soils. The hypothesis that ectomycorrhizal fungi on roots fractionate nitrogen isotopes leading to lower δ¹⁵N in needles is supported. This helps explain restoration outcomes, and bridges the gap between field and in vitro investigations.

     

Synergistic and antagonistic effects of mixing monospecific soils on plant-soil feedbacks

Background and aims

Disturbance affects the ability of organisms to persist on a site, and disturbance history acts as a filter of community composition. This is true for vascular plants and morphological groups of biocrusts, which respond differently to disturbance. Although functioning arid ecosystems include both groups, filtering of morphological groups of biocrusts has not previously been compared simultaneously with the responses of vascular plants.

Methods

Using a chronosequence approach, cover of vascular plants and biocrusts was examined across chronic disturbance gradients related to invasion by exotic species and grazing by livestock, following the acute disturbance of fire using paired burned and unburned plots in Wyoming big sagebrush on 99 plots.

Results

Cover of vascular plants and biocrusts was related to disturbance more so than abiotic factors of precipitation following fire, soil chemistry, percent coarse fragment and heat load index. Over time since fire of 12–23 years, we saw recovery of early successional groups: short mosses, shallow-rooted perennial grasses and annual forbs. Cover of deep and shallow-rooted perennial grasses and annual forbs increased in cover with intermediate levels of disturbance. Perennial forbs lacked a clear relationship with disturbance. Biocrusts decreased in cover with less disturbance when compared with perennial herbaceous plants but differed in sensitivities. Tall mosses were less sensitive to disturbance compared with lichens. Short mosses increased with some disturbance.

Conclusions

Morphological groups of biocrusts and vascular plants are eliminated with increasing variability in the size of gaps between perennials represented by the standard deviation of gaps between perennials. The inclusion of both groups in assessments of ecosystem recovery following disturbance addresses the fact that recovery of either group does not happen in isolation from the other but with interacting contributions to ecosystem functions.

     

Changes from pasture to a native tree plantation affect soil organic matter in a tropical soil,Panamá

Background and aims

We examined changes in soil organic matter arising from conversion of a 45-year old pasture to a 10 yr. old native tree plantation in Panamá, to evaluate the effect of monoculture and mixtures.

Methods

We intensively sampled the soil 0–10 cm depth in the pasture in 2001 and in 22 plantation plots in 2011, ranging from 5 monocultures to 3- and 6-species treatments; samples were also taken from an undisturbed forest site. Soil analyses included organic carbon (SOC) and δ¹³C.

Results

Conversion of the pasture to tree plantation resulted in an overall loss of SOC of 0.6 kg m^?2 (18%) in the top 10 cm, but neither tree species nor diversity had a significant effect. End-member δ¹³C values suggested that the contribution of C₃ plants to SOC was increased from 26% in the pasture to 55% after 10 years of plantation and SOC turnover times were calculated to be 21–36 yr.

Conclusions

The magnitude of the loss in soil SOC is smaller than the increases in tree biomass (~3 kg C m^?2) and litter (~0.3 kg C m^?2) in the plantation, but still a significant part of the ecosystem C balance.

     

Root architecture governs plasticity in response to drought

Aims

Fine root morphological traits are generally changed under soil nitrogen (N) enrichment, however, the underlying mechanism and functional significance are still not well understood. Our aims were to investigate the linkage of root morphology to anatomy, and its implication for root function at elevated soil N availability.

Methods

Ingrowth cores were used to sample root tips (0–20 cm soil depth) from six temperate tree species growing in monoculture plantations at a common site in northeastern China. Root morphological and anatomical traits were concurrently measured, and their relationships were determined within and among species in both control and N fertilization (10 g N m^?2y^?1) plots.

Results

Root diameter generally increased in all six species (non-significant for Phellodendron amurense) following N fertilization, which was caused by the increased root stele radius rather than cortical thickness. Congruently, N fertilization significantly decreased the ratio of cortical thickness to stele radius, but increased the ratio of total cross-sectional area of conduits to stele area in root tips across all species.

Conclusions

The observed anatomical changes of root tips contributed to the alternations of morphological root traits following N fertilization, with potentially important impacts on root physiological functions, like increased water and nutrient transport.

     

<Emphasis Type="Italic">massPix</Emphasis>: an R package for annotation and interpretation of mass spectrometry imaging data for lipidomics

Introduction

Mass spectrometry imaging (MSI) experiments result in complex multi-dimensional datasets, which require specialist data analysis tools.

Objectives

We have developed massPix—an R package for analysing and interpreting data from MSI of lipids in tissue.

Methods

massPix produces single ion images, performs multivariate statistics and provides putative lipid annotations based on accurate mass matching against generated lipid libraries.

Results

Classification of tissue regions with high spectral similarly can be carried out by principal components analysis (PCA) or k-means clustering.

Conclusion

massPix is an open-source tool for the analysis and statistical interpretation of MSI data, and is particularly useful for lipidomics applications.

     

Revegetation modifies patterns of temporal soil respiration responses to extreme-drying-and-rewetting in a semiarid ecosystem

Aims

The artificial cultivation of biocrusts may represent a new low-cost and highly efficient solution to erosion control. However, establishment under varying field environmental conditions is understudied. We tested a variety of methods, arriving at a set of technical recommendations for rapid establishment of moss biocrusts on disturbed slopes, and the industrialization of this process.

Methods

In multiple field experiments, aimed at moss biocrust cultivation and establishment, we considered the following factors: nutrient solutions (control and weekly addition); water-retaining agent (control and addition); plant growth regulator (control and biweekly addition); shading (0, 50%, 70% and 90%); dispersal method (broadcast and spray application). In all cases, we initially inoculated soils with 700 g/m² of moss biocrust materials. We monitored dynamic changes of the coverage and density of moss biocrusts during the cultivation period, and their biomass at the end.

Results

We successfully cultured moss biocrusts in a field setting in as little as two months. Specifically, we found:(1) Regardless of the dispersal method, the nutrient solutions and some degree of shading both increased the coverage, plant density and biomass of moss biocrusts, whereas the water-retaining agent and plant growth regulator had little influence on these parameters. The shading treatments improved the survival rates of moss biocrusts, with the shade rating of 70% exhibiting the best performance. Further, the nutrient solutions had a more positive effect under shaded conditions. (2) The growth of mosses dispersed in the fall exceeded that of mosses dispersed in the summer. (3) Under both dispersal techniques, the maximal coverage of the moss biocrusts exceeded 90%, and the maximal plant density of moss biocrusts reached 120 stems/cm²under broadcast dispersal, and 150 stems/cm², under spray dispersal.

Conclusions

The rapid restoration of moss biocrusts can best be achieved by spray-dispersal or broadcast-dispersal, while also applying Hoagland solution to supply nutrients and maintaining soil moisture at 15–25%. Fall inoculation appears more likely to lead to better moss establishment, in fact, high moss mortality occurred in summer unless shading was used. We have some evidence, observational in fall, and experimental in summer, that moderate shading favors establishment. This technique could feasibly be up scaled and adopted to restore some ecological functions on various types of engineered disturbed surfaces. Over a longer period, the survivorship, succession and sustainability of artificial moss biocrusts should be explored specifically.

     

Stable isotope signatures reflect dietary diversity in European forest moths

Background

Information on larval diet of many holometabolous insects remains incomplete. Carbon (C) and nitrogen (N) stable isotope analysis in adult wing tissue can provide an efficient tool to infer such trophic relationships. The present study examines whether moth feeding guild affiliations taken from literature are reflected in isotopic signatures.

Results

Non-metric multidimensional scaling and permutational analysis of variance indicate that centroids of dietary groups differ significantly. In particular, species whose larvae feed on mosses or aquatic plants deviated from those that consumed vascular land plants. Moth δ¹⁵N signatures spanned a broader range, and were less dependent on species identity than δ¹³C values. Comparison between moth samples and ostensible food sources revealed heterogeneity in the lichenivorous guild, indicating only Lithosia quadra as an obligate lichen feeder. Among root-feeding Agrotis segetum, some specimens appear to have developed on crop plants in forest-adjacent farm land. Reed-feeding stem-borers may partially rely on intermediary trophic levels such as fungal or bacterial growth.

Conclusion

Diagnostic partitioning of moth dietary guilds based on isotopic signatures alone could not be achieved, but hypotheses on trophic relationships based on often vague literature records could be assessed with high resolution. Hence, the approach is well suited for basic categorization of moths where diet is unknown or notoriously difficult to observe (i.e. Microlepidoptera, lichen-feeders).

     

Measurement of the <Superscript>15</Superscript>N/<Superscript>14</Superscript>N ratio of phenylalanine in fermentation matrix by isotope ratio mass spectrometry

Objectives

To determine the origin of ¹⁵N-labeled phenylalanine in microbial metabolic flux analysis using ¹⁵N as a tracer, a method for measuring phenylalanine δ¹⁵N using HPLC coupled with elemental analysis-isotope ratio mass spectrometry (EA-IRMS) was developed.

Results

The original source of the ¹⁵N-labeled phenylalanine was determined using this new method that consists of three steps: optimization of the HPLC conditions, evaluation of the isotope fractionation effects, and evaluation of the effect of pre-processing on the phenylalanine nitrogen stable isotope. In addition, the use of a ¹⁵N-labeled inorganic nitrogen source, rather than ¹⁵N-labeled amino acids, was explored using this method.

Conclusions

The method described here can also be applied to the analysis of metabolic flux.

     

Extraction of natural products from bark of <Emphasis Type="Italic">Betula pendula</Emphasis> using ionic liquids

Introduction

Aqueous–methanol mixtures have successfully been applied to extract a broad range of metabolites from plant tissue. However, a certain amount of material remains insoluble.

Objectives

To enlarge the metabolic compendium, two ionic liquids were selected to extract the methanol insoluble part of trunk from Betula pendula.

Methods

The extracted compounds were analyzed by LC/MS and GC/MS.

Results

The results show that 1-butyl-3-methylimidazolium acetate (IL-Ac) predominantly resulted in fatty acids, whereas 1-ethyl-3-methylimidazolium tosylate (IL-Tos) mostly yielded phenolic structures. Interestingly, bark yielded more ionic liquid soluble metabolites compared to interior wood.

Conclusion

From this one can conclude that the application of ionic liquids may expand the metabolic snapshot.

     

Characterization of ankylosing spondylitis and rheumatoid arthritis using <Superscript>1</Superscript>H NMR-based metabolomics of human fecal extracts

Introduction

The differences in fecal metabolome between ankylosing spondylitis (AS)/rheumatoid arthritis (RA) patients and healthy individuals could be the reason for an autoimmune disorder.

Objectives

The study explored the fecal metabolome difference between AS/RA patients and healthy controls to clarify human immune disturbance.

Methods

Fecal samples from 109 individuals (healthy controls 34, AS 40, and RA 35) were analyzed by ¹H NMR spectroscopy. Data were analyzed with principal component analysis (PCA) and orthogonal projection to latent structure discriminant (OPLS-DA) analysis.

Results

Significant differences in the fecal metabolic profiles could distinguish AS/RA patients from healthy controls but could not distinguish between AS and RA patients. The significantly decreased metabolites in AS/RA patients were butyrate, propionate, methionine, and hypoxanthine. Significantly increased metabolites in AS/RA patients were taurine, methanol, fumarate, and tryptophan.

Conclusion

The metabolome variations in feces indicated AS and RA were two homologous diseases that could not be distinguished by ¹H NMR metabolomics.

     

Acid and calcareous soils affect nitrogen nutrition and organic nitrogen uptake by beech seedlings (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) under drought,and their ectomycorrhizal community structure

Aims

The role of different soil types for beech productivity and drought sensitivity is unknown. The aim of this experimental study was to compare mycorrhizal diversity between acid sandy and calcareous soils and to investigate how this diversity affects tree performance, nitrogen uptake and use efficiency (NUE).

Methods

Beech trees were germinated and grown in five different soil types (pH 3.8 to 6.7). One-and-a-half-year-old plants were exposed for 6 weeks to sufficient or low soil humidity. Tree biomass, root tip mycorrhizal colonization and community structure, root tip mortality, leaf area, photosynthesis, nitrogen concentrations, NUE and short-term ¹⁵N uptake from glutamine were determined.

Results

Soil type did not affect photosynthesis or biomass formation, with one exception in calcareous soil, where root mortality was higher than in the other soil types. Beech in acid soils showed lower mycorrhizal colonization, higher nitrogen tissue concentrations, and lower NUE than those in calcareous soils. Drought had no effect on nitrogen concentrations or NUE but caused reductions in mycorrhizal colonization. Mycorrhizal species richness correlated with nitrogen uptake and NUE. Nitrogen uptake was more sensitive to drought in calcareous soils than in acid soils.

Conclusions

Beech may be more drought-susceptible on calcareous sites because of stronger decrease of organic nitrogen uptake than on acid soils.