Effects of litters with different concentrations of phenolics on the competition between Calluna vulgaris and Deschampsia flexuosa

We hypothesized that the outcome of competition between ericaceous plants and grasses is strongly affected by the concentrations of phenolics in the litter that they produce. To test the effect of phenolic-rich litter on soluble soil nitrogen concentrations, plant nitrogen uptake and inter-specific competition, we conducted a greenhouse experiment with the shrub Calluna vulgaris and the grass Deschampsia flexuosa and their leaf litters. Two litters of C. vulgaris were used, with equal nitrogen concentration but different (high and low) concentrations of total phenolics. The D. flexuosa leaf litter contained lower concentrations of phenolics, but higher concentrations of nitrogen than the C. vulgaris litters. The plants were grown in monocultures and in mixed cultures. Inorganic and dissolved organic nitrogen were measured monthly during the experiment. After four months, we measured above- and belowground biomass and the nutrient concentrations in above- and belowground plant parts. In monocultures, C. vulgaris produced more shoot and root biomass on its own litter than with no litter. Growth of Calluna was reduced on grass litter. D. flexuosa plants produced most biomass on their own litter type, whether in monocultures or in mixed cultures. Addition of Calluna litter stimulated the growth of D. flexuosa both in monoculture and in mixtures. The grass plants outcompeted Calluna both on shrub litter and on grass litter but not when grown without litter. The two C. vulgaris litter types that differed in their concentration of phenolics did not differ in their effects on the competition between the two species or on the production of inorganic and dissolved organic nitrogen. We conclude that the nitrogen content of the litter is more important as a plant feature driving competition between shrubs and grasses than the concentrations of phenolics.     

Estimating plant competition coefficients and predicting community dynamics from non-destructive pin-point data: a case study with Calluna?vulgaris and Deschampsia?flexuosa

A method is proposed for estimating plant competition coefficients and predicting the dynamics of herb and grassland plant communities from non-destructive pin-point measurements. The method is applied to inter-specific competition in a natural heathland community with relatively few interacting species. The study shows that the dynamics of the heathland plant community may be thought of as essentially a two-species system of Calluna vulgaris and Deschampsia flexuosa. There were significant competitive interactions between C. vulgaris and D. flexuosa. D. flexuosa affected both the cover and compactness of C. vulgaris individuals as a function of the compactness the previous year, whereas C. vulgaris significantly affected only the compactness of D. flexuosa. There was a significant negative effect of drought on the compactness of both C. vulgaris and D. flexuosa individuals, whereas night warming had no significant effects on either species. The predicted long-term outcome of the competitive interaction between C. vulgaris and D. flexuosa was that of unstable equilibrium, where the more dominant of the two will outcompete the other. However, when both species are found at relatively high plant covers the two species are predicted to co-exist for a long time period relatively to the time scale of the ageing process of C. vulgaris. Direct analyses of the inter-specific competitive interactions in natural plant communities with non-destructive measurements can provide important new insight into the processes that determine the composition of plant communities.     

High Resilience in Heathland Plants to Changes in Temperature, Drought, and CO2 in Combination: Results from the CLIMAITE Experiment

Climate change scenarios predict simultaneously increase in temperature, altered precipitation patterns and elevated atmospheric CO₂ concentration, which will affect key ecosystem processes and plant growth and species interactions. In a large-scale experiment, we investigated the effects of in situ exposure to elevated atmospheric CO₂ concentration, increased temperature and prolonged drought periods on the plant biomass in a dry heathland (Brandbjerg, Denmark). Results after 3 years showed that drought reduced the growth of the two dominant species Deschampsia flexuosa and Calluna vulgaris. However, both species recovered quickly after rewetting and the drought had no significant effect on annual aboveground biomass production. We did not observe any effects of increased temperature. Elevated CO₂ stimulated the biomass production for D. flexuosa in one out of three years but did not influence the standing biomass for either D. flexuosa or the ecosystem as more litter was produced. Treatment combinations showed little interactions on the measured parameters and in particular elevated CO₂ did not counterbalance the drought effect on plant growth, as we had anticipated. The plant community did not show any significant responses to the imposed climate changes and we conclude that the two heathland species, on a short time scale, will be relatively resistant to the changes in climatic conditions.     

Phenotypic plasticity in response to nitrate supply of an inherently fast-growing species from a fertile habitat and an inherently slow-growing species from an infertile habitat

The aim of the present study was to investigate possible differences in plasticity between a potentially fast-growing and a potentially slow-growing grass species. To this end, Holcus lanatus (L.) and Deschampsia flexuosa (L.) Trin., associated with fertile and infertile habitats, respectively, were grown in sand at eight nitrate concentrations. When plants obtained a fresh weight of approximately 5 g, biomass allocation, specific leaf area, the rate of net photosynthesis, the organic nitrogen concentration of various plant parts and the root weight at different soil depths were determined. There were linear relationships between the morphological and physiological features studied and the In-transformed nitrate concentration supplied, except for the specific leaf area and root nitrogen concentration of H. lanatus, which did not respond to the nitrate concentration. The root biomass of H. lanatus was invariably distributed over the soil layers than that of D. flexuosa. However, D. flexuosa allocated more root biomass to lower soil depths with decreasing nitrate concentration, in contrast to H. lanatus, which did not respond. The relative response to nitrate supply, i.e. the value of a character at a certain nitrate level relative to the value of that character at the highest nitrate supply, was used as a measure for plasticity. For a number of parameters (leaf area ratio, root weight ratio, root nitrogen concentration, vertical root biomass distribution and rate of net photosynthesis per unit leaf weight) the potentially slow-growing D. flexuosa exhibited a higher phenotypic plasticity than the potentially fast-growing H. lanatus. These findings are in disagreement with current literature. Possible explanations for this discrepancy are discussed in terms of differences in experimental approach as well as fundamental differences in specific traits between fast- and slow-growing grasses.     

Latent soil effects of grazing and ammonium deposition on Deschampsia flexuosa tillers inserted and grown in heather moorland soil

Upland heather moorlands support a range of increasingly rare and threatened biota, making them a priority habitat for conservation and restoration. Over-grazing and nitrogen deposition are two of the most important threats to maintaining these heather moorlands, yet there remains a paucity of research into their combined long-term effects. During the summer of 2008, we established an experiment within an existing research site at Pwllpeiran, mid-Wales. The original site, established in 1996, investigated long-term grazing and N deposition treatments (ammonium and nitrate) on heather moorland. Previous findings from the site, following a decade of treatment, suggested no significant, detectable impacts of treatments on soil C:N ratio and the foliar nitrogen of vegetation.The aim of our study was to investigate short- (2008) and long-term (1996–2008) N deposition treatment impacts, in combination with past grazing (1990–2007), on soil nutrient bioavailability. Soil cores were harvested and aboveground vegetation removed. Tillers of the grass Deschampsia flexuosa were planted into these cores which were then reciprocally transplanted back into the field experiment. The N deposition treatment was continued but grazing was excluded. D. flexuosa biomass changes were next assessed and leaf chemistry investigated using the metabolic fingerprinting method Fourier-transform infrared spectroscopy (FTIR) following three months of growth in the field (May–August 2008).Grazing treatment (on its own) had significant negative impacts on aboveground biomass and significant changes in plant chemistry were also revealed through the metabolic fingerprinting method Fourier transform infrared spectroscopy (FTIR). Short-term N deposition treatments during 2008 had no impacts on D. flexuosa growth or chemistry. There were also no detectable latent effects of long-term nitrate treatments on either growth or chemistry of D. flexuosa. However, plants grown in plots that had received long-term treatments of ammonium (NH₄⁺) had significantly lower poly-phenolic contents (revealed by FTIR) than plants grown in either nitrate (NO₃⁻) or control plots, suggesting detectable latent effects of N application in its reduced form. Further work needs to be undertaken to assess the relevance of residual soil nitrogen pools post N deposition and grazing.     

Nitrogen deposition and drought events have non-additive effects on plant growth – Evidence from greenhouse experiments

Climate change and nitrogen deposition affect biodiversity and ecosystem functioning, but interactive effects of these global change drivers are poorly understood. We analysed single and interactive effects of nitrogen (N) fertilisation and drought on the growth performance of Calluna vulgaris. We measured biomass production and allocation, tissue nutrient (N, phosphorus (P) and carbon (C)) concentrations, N allocation patterns (using ¹⁵N tracer) and plant's water status (using δ ¹³C signatures) as response variables in a 2-year greenhouse experiment. N fertilisation increased biomass production and biomass shoot:root ratios. ¹⁵N allocation patterns indicated an increasing aboveground N allocation following N fertilisation. Tissue δ ¹³C signatures were higher in N-fertilised plants. Plant responses to drought were weak. We found strong antagonistic interaction effects of N fertilisation and drought for biomass production. δ ¹³C values peaked when N-fertilised plants were subjected to drought, indicating that N fertilisation increased the evaporative demands of Calluna plants, likely due to increased biomass shoot:root ratios, which in turn resulted in higher drought susceptibility. As an important consequence, even slight drought events may weaken the competitiveness of Calluna when interacting with enhanced airborne N loads. Single-factor studies, thus, need to be complemented by multi-factor analyses to assess conceivable impacts of co-occurring global change drivers.     

Experimental manipulation of succession in heathland ecosystems

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession.     

Proteomics reveals the adaptability mechanism of Brassica napus to short-term boron deprivation

The application of pyrogenic carbon, biochar, to agricultural soils is currently discussed as a win-win strategy to sequester carbon in soil, thus improving soil fertility and mitigate global warming. Our aim was to investigate if biochar may improve plant eco-physiological responses under sufficient water supply as well as moderate drought stress. A fully randomized greenhouse study was conducted with the pseudo-cereal Chenopodium quinoa Willd, using three levels of biochar addition (0, 100 and 200?t ha^?1) to a sandy soil and two water treatments (60% and 20% of the water holding capacity of the control), investigating growth, water use efficiency, eco-physiological parameters and greenhouse gas (GHG) fluxes. Biochar application increased growth, drought tolerance and leaf-N- and water-use efficiency of quinoa despite larger plant?Cleaf areas. The plants growing in biochar-amended soil accumulated exactly the same amount of nitrogen in their larger leaf biomass than the control plants, causing significantly decreased leaf N-, proline- and chlorophyll-concentrations. In this regard, plant responses to biochar closely resembled those to elevated CO₂. However, neither soil- nor plant?Csoil-respiration was higher in the larger plants, indicating less respiratory C losses per unit of biomass produced. Soil-N₂O emissions were significantly reduced with biochar. The large application rate of 200?t ha^?1 biochar did not improve plant growth compared to 100?t ha^?1; hence an upper beneficial level exists. For quinoa grown in a sandy soil, biochar application might hence provide a win-win strategy for increased crop production, GHG emission mitigation and soil C sequestration.     

Nitrogen acquisition from inorganic and organic sources by boreal forest plants in the field

A wide range of recent studies have indicated that organic nitrogen may be of great importance to plant nitrogen (N) nutrition. Most of these studies have, however, been conducted in laboratory settings, excluding important factors for actual plant uptake, such as competition, mycorrhizal associations and soil interactions. In order to accurately evaluate the importance of different N compounds to plant N nutrition, field studies are crucial. In this study, we investigated short- as well as long-term plant nitrogen uptake by Deschampsia flexuosa, Picea abies and Vaccinium myrtillus from ¹⁵NO₃^–, ¹⁵NH₄⁺ and (U-¹³C, ¹⁵N) arginine, glycine or peptides. Root N uptake was analysed after 6 h and 64 days following injections. Our results show that all three species, irrespective of their type of associated mycorrhiza (arbuscular, ecto- or ericoid, respectively) rapidly acquired similar amounts of N from the entire range of added N sources. After 64 days, P. abies and V. myrtillus had acquired similar amounts of N from all N sources, while for D. flexuosa, the uptake from all N sources except ammonium was significantly lower than that from nitrate. Furthermore, soil analyses indicate that glycine was rapidly decarboxylated after injections, while other organic compounds exhibited slower turnover. In all, these results suggest that a wide range of N compounds may be of importance for the N nutrition of these boreal forest plants, and that the type of mycorrhiza may be of great importance for N scavenging, but less important to the N uptake capacity of plants.     

The importance of plant–soil interactions,soil nutrients,and plant life history traits for the temporal dynamics of Jacobaea vulgaris in a chronosequence of old‐fields

We examined to what extent temporal dynamics of Jacobaea vulgaris cover in old‐fields were related to plant–soil feedback, soil nutrients, seed availability and performance, and seedling establishment. Long‐term measurements at an experimental field and in ten old‐fields representing a chronosequence following land abandonment revealed a remarkably similar hump‐shaped temporal pattern of J. vulgaris cover, which peaked at about five years after abandonment. In a plant–soil feedback study, J. vulgaris biomass of plants grown in soil from all chronosequence fields was lower than in sterilized control soil. However, biomass of J. vulgaris in the feedback study was lower when grown in soil collected from fields with a high density of J. vulgaris plants than in soil from fields with a low density of J. vulgaris. When plants were grown again in the conditioned soil, a strong negative plant–soil feedback response was observed for soils from all fields. These results indicate that soils from all stages of the chronosequence can develop a strong negative soil feedback to J. vulgaris, and that there is a positive relationship between J. vulgaris density and the subsequent level of control by the soil community. In a common‐garden experiment with turfs collected from the chronosequence fields in which J. vulgaris was seeded, seedling establishment was significantly lower in turfs from older than from young fields. In a seed bank study the number of emerging seedlings declined with time since abandonment of the field. In conclusion, negative plant–soil feedback is an important factor explaining the hump‐shaped population development of J. vulgaris. However, it is not operating alone, as propagule availability and characteristics, and competition may also be important. Thus, in order to explain its contribution to plant population dynamics, the role of biotic plant–soil interactions, soil nutrients and life history characteristics along successional gradients should be considered from a community perspective.     

Effects of soil organic matter and nitrogen supply on competition between Festuca ovina and Deschampsia flexuosa during inland dune succession

We tested the prediction that the successional replacement of plant species during succession on inland sand dunes results from the effects of an increase in nitrogen mineralization on competitive interactions. The growth and competitive strength of Festuca ovina and Deschampsia flexuosa on soil substrates with different amounts of soil organic matter or nitrogen supply were measured. Small tillers of Festuca ovina and Deschampsia flexuosa were grown in monocultures and 1:1-mixtures on soil columns with undisturbed layers of soil organic matter from different successional age. There was (a) no visible soil organic matter, (b) a thin soil organic layer (0.5 cm) and (c) a thicker soil organic layer (6.0 cm) present on the soil columns. The species were also grown on columns with no visible soil organic matter (bare sand) with two different levels of N fertilization to mimic the increased N mineralization in the older successional stages.In monoculture, Festuca produced more biomass on the substrates with a soil organic layer compared to the unfertilized sand substrate. It also produced more biomass on sand substrates with N fertilization. Deschampsia produced more biomass in treatments with a soil organic layer compared to the bare sand treatments, but did not respond to the ammonium-nitrate addition. In competition, Festuca seemed to be the stronger competitor on the unfertilized sand substrate. Festuca was also the better competitor on the N fertilized sand treatments, while on the treatments with a soil organic layer Deschampsia was the winning species. Our results do not support the hypothesis that an increase in N supply is responsible for the replacement of Festuca by Deschampsia that concur with the accumulation of soil organic matter during succession in inland dunes.     

Biosynthesis of plant phenolic compounds in elevated atmospheric CO2

Experiments were carried out to determine the effects of elevated atmospheric carbon dioxide (CO₂) on phenolic biosynthesis in four plant species growing over three generations for nine months in a model plant community. Results were compared to those obtained when the same species were grown individually in pots in the same soils and controlled environment. In the model herbaceous plant community, only two of the four species showed any increase in biomass under elevated CO₂, but this occurred only in the first generation for Spergula arvensis and in the second generation for Poa annua. Thus, the effects of CO₂ on plant biomass and carbon and nitrogen content were species‐ and generation‐specific. The activity of the principle phenolic biosynthetic enzyme, phenylalanine ammonia lyase (PAL), increased under elevated CO₂ in Senecio vulgaris only in Generation 1, but increased in three of the four plant species in Generation 2. There were no changes in the total phenolic content of the plants, except for P. annua in Generation 1. Lignin content decreased under elevated CO₂ in Cardamine hirsuta in Generation 1, but increased in Generation 2, whilst the lignin content of P. annua showed no change, decreased, then increased in response to elevated CO₂ over the three generations. When the species were grown alone in pots, elevated CO₂ increased PAL activity in plants grown in soil taken from the Ecotron community after nine months of plant growth, but not in plants grown in the soil used at the start of the experiment (‘initial' soil). In P. annua, phenolic biosynthesis decreased under elevated CO₂ in initial soil, and in both P. annua and S. vulgaris there was a significant interaction between effects of soil type and CO₂ level on PAL activity. In this study, plant chemical composition altered more in response to environmental factors such as soil type than in response to carbon supply. Results were species‐specific and changed markedly between generations.     

Effect of drought and low light on growth and enzymatic antioxidant system of Picea asperata seedlings

The combined effects of drought and low light on biomass partition, foliar nitrogen concentration, membrane stability and active oxygen species (AOS) and antioxidant system were investigated in dragon spruce (Picea asperata Mast.) seedlings grown at two watering regimes (well-watered, 100% of field capacity and drought, 30% of field capacity) and light availabilities (HL, 100% of full sunlight and low light, 15% of full sunlight). Under high light condition drought not only reduced foliar nitrogen concentration (Nmass) and membrane stability index (MSI) but also significantly increased biomass partitioning to roots, AOS, ascorbic acid (AsA) content and antioxidant enzyme activities including superoxide dismutase (SOD, EC 1.15.1.1), peroxidase (POD, EC 1.11.1.7), catalase (CAT, EC 1.11.1.6), ascorbate peroxidase (APX, EC 1.11.1.11) and glutathione reductase(GR, EC 1.6.4.2). However, no prominently drought-induced differences in biomass partitioning to root, SOD, GR activities, hydrogen peroxide (H₂O₂) and MSI were observed in low light seedlings. On the other hand, significant interaction of drought and low light was found on MSI, the antioxidant enzymes activities (SOD, POD, CAT, APX, GR), H₂O₂ and superoxide radical (O₂ ⁻). These results suggested that seedlings grown at the understory were more sensitive to drought than low light.     

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.

     

Off-season uptake of nitrogen in temperate heath vegetation

In this field study we show that temperate coastal heath vegetation has a significant off-season uptake potential for nitrogen, both in the form of ammonium and as glycine, throughout winter. We injected ¹⁵N-ammonium and ¹⁵N 2×(¹³C)-glycine into the soil twice during winter and once at spring. The winter temperatures were similar to those of an average winter in the northern temperate region of Europe, with only few days of soil temperatures below zero or above 5°C. The vegetation, consisting of the evergreen dwarf shrub Calluna vulgaris, the deciduous dwarf shrub Salix arenaria, and the graminoids Carex arenaria and Deschampsia flexuosa, showed high root uptake of both forms of nitrogen, both 1 day after labelling and after a month, in species specific temporal patterns. Plant uptake of ¹³C was not significant, providing no further evidence of intact uptake of glycine. Translocation of the labelled nitrogen to shoots was generally evident after 1 month and increased as spring approached, with different translocation strategies in the three plant functional types. Furthermore, only the graminoids showed shoot growth during winter. Increasing plant nitrogen concentration from fall to spring at temperate heaths may, hence, be due to nitrogen uptake. Our results suggest that the potential for nitrogen uptake in plants at winter is of the same order of magnitude as at summer. Hence, winter nitrogen uptake in ecosystems in the temperate/boreal region should be considered when making annual nitrogen budgets of heath ecosystems, and the view of plant nutrient uptake as low in this climatic region during winter should be revised.     

Influence of water availability on competitive interactions between plant species on sandy soils

The gravimetric soil water content of four different successional stages of inland dune vegetation revealed pronounced seasonal changes and particularly a major drought period in the late summer. In order to analyze the effect of these seasonal water changes on the competitive interactions of two dominant species (Carex arenaria andDeschampsia flexuosa) in one of these stages root development was compared under experimentally manipulated water conditions in a minirhizotron experiment. Under well-watered conditions, the root growth ofC. arenaria andD. flexuosa was not influenced by competition, but was significantly reduced forC. arenaria at low water availability. Accordingly, the relative yield based on total biomass was significantly lower under water shortage than under well-watered conditions forC. arenaria, indicating higher interspecific competition compared to intraspecific competitive interactions. These results indicate (i) that soil water status is an important factor for belowground competitive interactions and (ii) that seasonal changes in soil water content may have a pronounced effect on the outcome of competition and, in consequence, on the structure of the respective plant communities.     

Energy Demands of Nitrogen Supply in Mass Cultivation of Two Commercially Important Microalgal Species, <Emphasis Type="Italic">Chlorella vulgaris</Emphasis> and <Emphasis Type="Italic">Dunaliella tertiolecta</Emphasis>

Mass culture of microalgae is a potential alternative to cultivation of terrestrial crops for bioenergy production. However, microalgae require nitrogen fertiliser in quantities much higher than plants, and this has important consequences for the energy balance of these systems. The effect of nitrogen fertiliser supplied to microalgal bubble-column photobioreactor cultures was investigated using different nitrogen sources (nitrate, urea, ammonium) and culture conditions (air, 12% CO₂). In 20 L cultivations, maximum biomass productivity for Chlorella vulgaris cultivated using nitrate and urea was 0.046 and 0.053 g L⁻¹ day⁻¹, respectively. Maximum biomass productivity for Dunaliella tertiolecta cultivated using nitrate, urea and ammonium was 0.033, 0.038 and 0.038 g L⁻¹ day⁻¹, respectively. In intensive bubble-column photobioreactors using 12% CO₂, maximum productivity reached 0.60 and 0.83 g L⁻¹ day⁻¹ for C. vulgaris and D. tertiolecta, respectively. Recycling of nitrogen within the photobioreactor system via algal exudation of nitrogenous compounds and bacterial activity was identified as a potentially important process. The energetic penalty incurred by supply of artificial nitrogen fertilisers, phosphorus, power and CO₂ to microalgal photobioreactors was investigated, although analysis of all energy burdens from biomass production to usable energy carriers was not conducted. After subtraction of the power, nitrogen and phosphorus energy burdens, maximum net energy ratios for C. vulgaris and D. tertiolecta cultivated in bubble columns were 1.82 and 2.10. Assuming CO₂ was also required from a manufactured source, the net energy ratio decreased to 0.09 and 0.11 for C. vulgaris and D. tertiolecta, so that biomass production in this scenario was unsustainable. Although supply of nitrogen is unlikely to be the most energetically costly factor in sparged photobioreactor designs, it is still a very significant penalty. There is a need to optimise both cultivation strategies and recycling of nitrogen in order to improve performance. Data are supported by measurements including biochemical properties (lipid, protein, heating value) and bacterial number by epifluorescence microscopy.     

Influence of arbuscular mycorrhizal fungi and simulated acid rain on the growth and coexistence of the grasses Calamagrostis villosa and Deschampsia flexuosa

Two grass species — Calamagrostis villosa (Chaix) J.F. Gmelin and Deschampsia flexuosa (L.) Trin. — are expanding in mountain Norway spruce (Picea abies L. Karst.) forests of Central Europe damaged by anthropogenic pollution constituted particularly of acid rain. This invasion of grasses may be caused by the higher irradiance reaching the forest floor after the pollution-induced tree defoliation. The relative abundance of the two grass species is changing during the process of forest decline. Our study investigated the effects of arbuscular mycorrhizal fungi (AMF) on the growth and coexistence of both species under simulated acid rain (SAR) and two levels of irradiance. Three microcosm experiments were conducted to investigate how both grasses are influenced by the AMF when grown separately or together interacting via extraradical mycelium (ERM). A positive growth response to inoculation with Glomus mosseae BEG 25 was found for both grass species when cultivated separately and the mycorrhizal dependence and the growth benefit for D. flexuosa was greater than for C. villosa. However, when both grass species were grown together in the rhizoboxes with separated root and hyphal compartments, the growth effect of the AMF was the opposite, i.e. C. villosa benefited more. The plants did not benefit from the AMF inoculation under the SAR treatment compared with dH2O treatment. The SAR also negatively influenced root length colonised by AMF, length of the ERM, alkaline phosphatase and NADH diaphorase activities of the ERM. The role of the ERM in transporting phosphorus between these grasses was verified by applying the radioisotope ³²P. There was a greater transport of isotopic ³²P between inoculated plants C. villosa and D. flexuosa grown in separated root compartments, as compared to non-inoculated plants. The amount of transported ³²P was low: a maximum of 3% of applied ³²P was detected in the shoots of receiver plants. Mechanical disturbance of the ERM significantly decreased the ³² P transport between plants. The ³²P transport between mycorrhizal plants was higher in the D. flexuosa to C. villosa direction than in the opposite one. Neither the SAR nor the low level of irradiance influenced the amount of transported ³²P. We discuss the role of ERM links between root systems in the coexistence of both grass species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Inability to solubilize phosphate in limestone soils—key factor controlling calcifuge habit of plants

Germination, seedling establishment and growth of calcifuge plants in Swedish limestone soils of Archean and Ordovician age were studied. As previously demonstrated for Viscaria vulgaris, establishment of Rumex acetosella and Silene rupestris did not succeed unless CaHPO₄ (at the rate of 10 mmol dm^-3 of soil) was supplied. Growth of Deschampsia flexuosa was enhanced by phosphate addition, whereas establishment success of Jasione montana was poor, regardless of phosphate treatment. Establishment and growth in an acidic gneiss soil, used as a reference for the species studied, was good. Total, total inorganic, exchangeable, and soil solution P were considered in all soils and treatments. It is proposed that the calcifuge behaviour of plants is quite often caused by inability to solubilize the native phosphate of limestone soils.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}