Petra Fransson

Elevated CO2 and nitrogen influence exudation of soluble organic compounds by ectomycorrhizal root systems

Root and mycelial exudation contributes significantly to soil carbon (C) fluxes, and is likely to be altered by an elevated atmospheric carbon dioxide (CO(2)) concentration and nitrogen (N) deposition. We quantified soluble, low-molecular-weight (LMW) organic compounds exuded by ectomycorrhizal plants grown under ambient (360 p.p.m.) or elevated (710 p.p.m.) CO(2) concentrations and with different N sources. Scots pine seedlings, colonized by one of five different ectomycorrhizal or nonmycorrhizal fungi, received 70 muM N, either as NH(4)Cl or as alanine, in a liquid growth medium. Exudation of LMW organic acids (LMWOAs), dissolved monosaccharides and total dissolved organic carbon were determined. Both N and CO(2) had a significant impact on exudation, especially of LMWOAs. Exudation of LMWOAs was negatively affected by inorganic N and decreased by 30-85% compared with the organic N treatment, irrespective of the CO(2) treatment. Elevated CO(2) had a clear impact on the production of individual LMWOAs, although with very contrasting effects depending on which N source was supplied.

Quantitative analysis of root and ectomycorrhizal exudates as a response to Pb, Cd and As stress

We examined exudation of low molecular weight (LMW) organic compounds of ectomycorrhizal (ECM) and non-mycorrhizal (NM) seedlings in relation to metals. Scots pine seedlings, either colonized by one of six different ECM fungi or NM, were grown in Petri dishes containing glass beads and liquid growth medium and exposed to elevated concentrations of Pb, Cd and As. Exudation of LMW organic compounds (LMW organic acids (LMWOAs), amino acids and dissolved monosaccharides) and dissolved organic carbon (DOC) was determined qualitatively and quantitatively and exudation rates were calculated. Metals had a significant impact on exudation, especially of oxalate. For Pb and Cd treatments, exudation of oxalate and total LMWOAs generally increased by 15–45% compared to nutrient controls. Production of amino acids, dissolved monosaccharides and DOC was not significantly stimulated by exposure to metals; however, there were non-significant trends towards increased exudation. Finally, exudation generally increased in the presence of mycorrhizal seedlings compared to NM seedlings. The results suggest that ECM fungi may reduce the toxicity of metals to plants through significant increases in the production of organic chelators. Axenic conditions are required to assess the full potential for production of these molecules but their overall significance in soil ecosystems needs to be determined using additional experiments under more ecologically realistic conditions.

Phosphorus cycling in deciduous forest soil differs between stands dominated by ecto- and arbuscular mycorrhizal trees

Although much is known about how trees and their associated microbes influence nitrogen cycling in temperate forest soils, less is known about biotic controls over phosphorus (P) cycling. Given that mycorrhizal fungi are instrumental for P acquisition and that the two dominant associations - arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi - possess different strategies for acquiring P, we hypothesized that P cycling would differ in stands dominated by trees associated with AM vs ECM fungi. We quantified soil solution P, microbial biomass P, and sequentially extracted inorganic and organic P pools from May to November in plots dominated by trees forming either AM or ECM associations in south-central Indiana, USA. Overall, fungal communities in AM and ECM plots were functionally different and soils exhibited fundamental differences in P cycling. Organic forms of P were more available in ECM plots than in AM plots. Yet inorganic P decreased and organic P accumulated over the growing season in both ECM and AM plots, resulting in increasingly P-limited microbial biomass. Collectively, our results suggest that P cycling in hardwood forests is strongly influenced by biotic processes in soil and that these are driven by plant-associated fungal communities.

Traits to Ecosystems: The Ecological Sustainability Challenge When Developing Future Energy Crops

Today we are undertaking great efforts to improve biomass production and quality traits of energy crops. Major motivation for developing those crops is based on environmental and ecological sustainability considerations, which however often are de-coupled from the trait-based crop improvement programs. It is now time to develop appropriate methods to link crop traits to production system characteristics set by the plant and the biotic communities influencing it; and to the ecosystem processes affecting ecological sustainability. The relevant ecosystem processes involve the net productivity in terms of biomass and energy yields, the depletion of energy-demanding resources (e.g. nitrogen, N), the carbon dynamics in soil and atmosphere, and the resilience and temporal stability of the production system. In a case study, we compared aspects of N use efficiency in various varieties of an annual (spring wheat) and perennial (Salix) energy crop grown under two nutrient regimes in Sweden. For example, we found considerable variation among crops, varieties and nutrient regimes in the energy yield per plant-internal N (MJ g-1 yr-1), which would result in different N resource depletion per unit energy produced.

Correspondence of ectomycorrhizal diversity and colonisation of willows (Salix spp.) grown in short rotation coppice on arable sites and adjacent natural stands.

Willows (Salix spp.) are mycorrhizal tree species sometimes cultivated as short rotation coppice (SRC) on arable sites for energy purposes; they are also among the earliest plants colonising primary successional sites in natural stands. The objective of this study was to analyse the degree of colonisation and diversity of ectomycorrhizal (EM) communities on willows grown as SRC in arable soils and their adjacent natural or naturalized stands. Arable sites usually lack ectomycorrhizal host plants before the establishment of SRC, and adjacent natural or naturalized willow stands were hypothesized to be a leading source of ectomycorrhizal inoculum for the SRC. Three test sites including SRC stands (Salix viminalis, Salix dasyclados, and Salix schwerinii) and adjacent natural or naturalized (Salix caprea, Salix fragilis, and Salix × mollissima) stands in central Sweden were investigated on EM colonisation and morphotypes, and the fungal partners of 36 of the total 49 EM fungi morphotypes were identified using molecular tools. The frequency of mycorrhizas in the natural/naturalized stands was higher (two sites) or lower (one site) than in the corresponding cultivated stands. Correspondence analysis revealed that some EM taxa (e.g. Agaricales) were mostly associated with cultivated willows, while others (e.g. Thelephorales) were mostly found in natural/naturalized stands. In conclusion, we found strong effects of sites and willow genotype on EM fungi formation, but poor correspondence between the EM fungi abundance and diversity in SRC and their adjacent natural/naturalized stands. The underlying mechanism might be selective promotion of some EM fungi species by more effective spore dispersal.

Host Plant—Ectomycorrhizal Fungus Combination Drives Resource Allocation in Willow: Evidence for Complex Species Interaction from a Simple Experiment

Abstract Root-associated mycorrhizal fungi affect plant growth and resource allocation. Our major aim was to explore the plant-internal mechanisms behind the effects of mycorrhizal colonization on leaf chemistry of willow (Salix spp.). Combinations of 2 willow varieties (Loden, Tora) and 2 ectomycorrhizal fungal species (Hebeloma fastibile, Tricholoma cingulatum) were grown under controlled conditions. Host plant variety and fungal species effects on host resource allocation (biomass and leaf chemistry) varied in a complex way. Shoot biomass growth and allocation was mostly affected by host plant variety, whereas leaf and root biomass allocation were strongly affected by mycorrhizal treatment. Leaf biomass production was affected by willow variety, mycorrhizal treatment, and the interaction between them. The results indicate a strong effect of mycorrhizal colonization on host plant biomass allocation, which can mediate mycorrhizal effects on leaf chemistry. For example, leaf biomass allocation was inversely correlated with foliar concentrations of salicylic acid, suggesting a functional link between the two. However, the apparent complexity of the host plant variety-fungal species interaction makes any prediction of their possible outcome very difficult, especially in an ecological context. Still, fungal species identity seems to be more important for the responses of Salix to mycorrhizal colonization than the amount of fungal biomass.

Albatrellus citrinus sp. nov., connected to Picea abies on lime rich soils

Field observations indicated that a morphotype of Albatrellus subrubescens seemed connected to Picea abies and lime rich soils, while the original morphotype seemed connected with Pinus sylvestris and indifferent to lime. We conducted a molecular study (ITS sequencing of 22 Albatrellus specimens) to test the hypothesis that we in fact had discovered a new species. Our results confirmed the hypothesis, i.e. the Picea taxon (604 bp ITS) had little intraspecific variation in spite of 1600 km distance between samples, but compared with the Pinus taxon (598 bp ITS) the sequence difference was constantly 5.6% regardless of close distance (75 km). We describe the new species A. citrinus, which apart from ecology and ITS sequence, is different from A. subrubescens by a distinct yellowing with age, lack of dark spots of the cap, a mild taste and somewhat narrower spores. A. citrinus seems to be more related to A. ovinus than to A. subrubescens, and A. syringae may not even be a true Albatrellus.

Two Salix Genotypes Differ in Productivity and Nitrogen Economy When Grown in Monoculture and Mixture

Individual plant species or genotypes often differ in their demand for nutrients; to compete in a community they must be able to acquire more nutrients (i.e., uptake efficiency) and/or use them more efficiently for biomass production than their competitors. These two mechanisms are often complementary, as there are inherent trade-offs between them. In a mixed-stand, species with contrasting nutrient use patterns interact and may use their resources to increase productivity in different ways. Under contrasting nutrient availabilities, the competitive advantages conferred by either strategy may also shift, so that the interaction between resource use strategy and resource availability ultimately determines the performance of individual genotypes in mixtures. The aim was to investigate growth and nitrogen (N) use efficiency of two willow (Salix) genotypes grown in monoculture and mixture in a fertilizer contrast. We explored the hypotheses that (1) the biomass production of at least one of the involved genotypes should be greater when grown in mixture as compared to the corresponding monoculture when nutrients are the most growth-limiting factor; and (2) the N economy of individual genotypes differs when grown in mixture compared to the corresponding monoculture. The genotypes ‘Tora’ (Salix schwerinii ×S. viminalis) and ‘Loden’ (S. dasyclados), with contrasting phenology and functional traits, were grown from cuttings in a growth container experiment under two nutrient fertilization treatments (high and low) in mono- and mixed-culture for 17 weeks. Under low nutrient level, ‘Tora’ showed a higher biomass production (aboveground biomass, leaf area productivity) and N uptake efficiency in mixture than in monoculture, whereas ‘Loden’ showed the opposite pattern. In addition, ‘Loden’ showed higher leaf N productivity but lower N uptake efficiency than ‘Tora.’ The results demonstrated that the specific functional trait combinations of individual genotypes affect their response to mixture as compared to monoculture. Plants grown in mixture as opposed to monoculture may thus increase biomass and vary in their response of N use efficiency traits. However, young plants were investigated here, and as we cannot predict mixture response in mature stands, our results need to be validated at field scale.

Mixture of Salix Genotypes Promotes Root Colonization With Dark Septate Endophytes and Changes P Cycling in the Mycorrhizosphere

The roots of Salix spp. can be colonized by two types of mycorrhizal fungi (ectomycorrhizal and arbuscular) and furthermore by dark-septate endophytes. The fungal root colonization is affected by the plant genotype, soil properties and their interactions. However, the impact of host diversity accomplished by mixing different Salix genotypes within the site on root-associated fungi and P-mobilization in the field is not known. It can be hypothesized that mixing of genotypes with strong eco-physiological differences changes the diversity and abundance of root-associated fungi and P-mobilization in the mycorrhizosphere based on different root characteristics. To test this hypothesis, we have studied the mixture of two fundamentally eco-physiologically different Salix genotypes (S. dasyclados cv. ‘Loden’ and S. schwerinii × S. viminalis cv. ‘Tora’) compared to plots with pure genotypes in a randomized block design in a field experiment in Northern Germany. We assessed the abundance of mycorrhizal colonization, fungal diversity, fine root density in the soil and activities of hydrolytic enzymes involved in P-mobilization in the mycorrhizosphere in autumn and following spring after three vegetation periods. Mycorrhizal and endophytic diversity was low under all Salix treatments with Laccaria tortilis being the dominating ectomyorrhizal fungal species, and Cadophora and Paraphaeosphaeria spp. being the most common endophytic fungi. Interspecific root competition increased richness and root colonization by endophytic fungi (four taxa in the mixture vs. one found in the pure host genotype cultures) more than by ectomycorrhizal fungi and increased the activities of hydrolytic soil enzymes involved in the P-mineralization (acid phosphatase and β-glucosidase) in mixed stands. The data suggest selective promotion of endophytic root colonization and changed competition for nutrients by mixture of Salix genotypes.