Directly quantifying multiple interacting influences on plant competition.

When plants compete what influences that interaction? To answer this we measured belowground competition directly, as the simultaneous capture of soil ammonium and nitrate by co-existing herbaceous perennials, Dactylis glomerata and Plantago lanceolata, under the influence of: species identity; N uptake and biomass of focal and neighbour plants; location (benign lowland versus harsher upland site); N availability (low or high N fertilizer); N ion, ammonium or nitrate production (mineralisation) rate, and competition type (intra- or interspecific), as direct effects or pairwise interactions in linear models. We also measured biomass as an indirect proxy for competition. Only three factors influenced both competitive N uptake and biomass production: focal species identity, N ion and the interaction between N ion and neighbour N uptake. Location had little effect on N uptake but a strong influence on biomass production. N uptake increased linearly with biomass only in isolated plants. Our results support the view that measuring resource capture or biomass production tells you different things about how competitors interact with one another and their environment, and that biomass is a longer-term integrative proxy for the outcomes of multiple separate interactions-such as competition for N-occurring between plants.

How are nitrogen availability, fine-root mass, and nitrogen uptake related empirically? Implications for models and theory.

Understanding the effects of global change in terrestrial communities requires an understanding of how limiting resources interact with plant traits to affect productivity. Here, we focus on nitrogen and ask whether plant community nitrogen uptake rate is determined (a) by nitrogen availability alone or (b) by the product of nitrogen availability and fine-root mass. Surprisingly, this is not empirically resolved. We performed controlled microcosm experiments and reanalyzed published pot experiments and field data to determine the relationship between community-level nitrogen uptake rate, nitrogen availability, and fine-root mass for 46 unique combinations of species, nitrogen levels, and growing conditions. We found that plant community nitrogen uptake rate was unaffected by fine-root mass in 63% of cases and saturated with fine-root mass in 29% of cases (92% in total). In contrast, plant community nitrogen uptake rate was clearly affected by nitrogen availability. The results support the idea that although plants may over-proliferate fine roots for individual-level competition, it comes without an increase in community-level nitrogen uptake. The results have implications for the mechanisms included in coupled carbon-nitrogen terrestrial biosphere models (CN-TBMs) and are consistent with CN-TBMs that operate above the individual scale and omit fine-root mass in equations of nitrogen uptake rate but inconsistent with the majority of CN-TBMs, which operate above the individual scale and include fine-root mass in equations of nitrogen uptake rate. For the much smaller number of CN-TBMs that explicitly model individual-based belowground competition for nitrogen, the results suggest that the relative (not absolute) fine-root mass of competing individuals should be included in the equations that determine individual-level nitrogen uptake rates. By providing empirical data to support the assumptions used in CN-TBMs, we put their global climate change predictions on firmer ground.

Long-term impacts of nitrogen deposition on coastal plant communities.

Nitrogen deposition has been shown to have significant impacts on a range of vegetation types resulting in eutrophication and species compositional change. Data from a re-survey of 89 coastal sites in Scotland, UK, c. 34 years after the initial survey were examined to assess the degree of change in species composition that could be accounted for by nitrogen deposition. There was an overall increase in the Ellenberg Indicator Value for nitrogen (EIV-N) of 0.15 between the surveys, with a clear shift to species characteristic of more eutrophic situations. This was most evident for Acid grassland, Fixed dune, Heath, Slack and Tall grass mire communities and despite falls in EIV-N for Improved grass, Strand and Wet grassland. The increase in EIV-N was highly correlated to the cumulative deposition between the surveys, and for sites in south-east Scotland, eutrophication impacts appear severe. Unlike other studies, there appears to have been no decline in species richness associated with nitrogen deposition, though losses of species were observed on sites with the very highest levels of SOx deposition. It appears that dune vegetation (specifically Fixed dune) shows evidence of eutrophication above 4.1 kg N ha(-1) yr(-1), or 5.92 kg N ha(-1) yr(-1) if the lower 95% confidence interval is used. Coastal vegetation appears highly sensitive to nitrogen deposition, and it is suggested that major changes could have occurred prior to the first survey in 1976.

Species composition of coastal dune vegetation in Scotland has proved resistant to climate change over a third of a century.

Climate change is expected to have an impact on plant communities as increased temperatures are expected to drive individual species' distributions polewards. The results of a revisitation study after c. 34 years of 89 coastal sites in Scotland, UK, were examined to assess the degree of shifts in species composition that could be accounted for by climate change. There was little evidence for either species retreat northwards or for plots to become more dominated by species with a more southern distribution. At a few sites where significant change occurred, the changes were accounted for by the invasion, or in one instance the removal, of woody species. Also, the vegetation types that showed the most sensitivity to change were all early successional types and changes were primarily the result of succession rather than climate-driven changes. Dune vegetation appears resistant to climate change impacts on the vegetation, either as the vegetation is inherently resistant to change, management prevents increased dominance of more southerly species or because of dispersal limitation to geographically isolated sites.

Dynamic trajectories of growth and nitrogen capture by competing plants.

Although dynamic, plant competition is usually estimated as biomass differences at a single, arbitrary time; resource capture is rarely measured. This restricted approach perpetuates uncertainty. To address this problem, we characterized the competitive dynamics of Dactylis glomerata and Plantago lanceolata as continuous trajectories of biomass production and nitrogen (N) capture. Plants were grown together or in isolation. Biomass and N content were measured at 17 harvests up to 76 d after sowing. Data were fitted to logistic models to derive instantaneous growth and N capture rates. Plantago lanceolata was initially more competitive in terms of cumulative growth and N capture, but D. glomerata was eventually superior. Neighbours reduced maximum biomass, but influenced both maximum N capture and its rate constant. Timings of maximal instantaneous growth and N capture rates were similar between species when they were isolated, but separated by 16 d when they were competing, corresponding to a temporal convergence in maximum growth and N capture rates in each species. Plants processed N and produced biomass differently when they competed. Biomass and N capture trajectories demonstrated that competitive outcomes depend crucially on when and how 'competition' is measured. This potentially compromises the interpretation of conventional competition experiments.

A new hammer to crack an old nut: interspecific competitive resource capture by plants is regulated by nutrient supply, not climate.

Although rarely acknowledged, our understanding of how competition is modulated by environmental drivers is severely hampered by our dependence on indirect measurements of outcomes, rather than the process of competition. To overcome this, we made direct measurements of plant competition for soil nitrogen (N). Using isotope pool-dilution, we examined the interactive effects of soil resource limitation and climatic severity between two common grassland species. Pool-dilution estimates the uptake of total N over a defined time period, rather than simply the uptake of ¹5N label, as used in most other tracer experiments. Competitive uptake of N was determined by its available form (NO3⁻ or NH4⁺). Soil N availability had a greater effect than the climatic conditions (location) under which plants grew. The results did not entirely support either of the main current theories relating the role of competition to environmental conditions. We found no evidence for Tilman's theory that competition for soil nutrients is stronger at low, compared with high nutrient levels and partial support for Grime's theory that competition for soil nutrients is greater under potentially more productive conditions. These results provide novel insights by demonstrating the dynamic nature of plant resource competition.

Root-shoot growth responses during interspecific competition quantified using allometric modelling.

BACKGROUND AND AIMS: Plant competition studies are restricted by the difficulty of quantifying root systems of competitors. Analyses are usually limited to above-ground traits. Here, a new approach to address this issue is reported. METHODS: Root system weights of competing plants can be estimated from: shoot weights of competitors; combined root weights of competitors; and slopes (scaling exponents, α) and intercepts (allometric coefficients, ß) of ln-regressions of root weight on shoot weight of isolated plants. If competition induces no change in root : shoot growth, α and ß values of competing and isolated plants will be equal. Measured combined root weight of competitors will equal that estimated allometrically from measured shoot weights of each competing plant. Combined root weights can be partitioned directly among competitors. If, as will be more usual, competition changes relative root and shoot growth, the competitors' combined root weight will not equal that estimated allometrically and cannot be partitioned directly. However, if the isolated-plant α and ß values are adjusted until the estimated combined root weight of competitors matches the measured combined root weight, the latter can be partitioned among competitors using their new α and ß values. The approach is illustrated using two herbaceous species, Dactylis glomerata and Plantago lanceolata. KEY RESULTS: Allometric modelling revealed a large and continuous increase in the root : shoot ratio by Dactylis, but not Plantago, during competition. This was associated with a superior whole-plant dry weight increase in Dactylis, which was ultimately 2·5-fold greater than that of Plantago. Whole-plant growth dominance of Dactylis over Plantago, as deduced from allometric modelling, occurred 14-24 d earlier than suggested by shoot data alone. CONCLUSION: Given reasonable assumptions, allometric modelling can analyse competitive interactions in any species mixture, and overcomes a long-standing problem in studies of competition.

Interactions among fungal community structure, litter decomposition and depth of water table in a cutover peatland.

Peatlands are important reservoirs of carbon (C) but our understanding of C cycling on cutover peatlands is limited. We investigated the decomposition over 18 months of five types of plant litter (Calluna vulgaris, Eriophorum angustifolium, Eriophorum vaginatum, Picea sitchensis and Sphagnum auriculatum) at a cutover peatland in Scotland, at three water tables. We measured changes in C, nitrogen (N) and phosphorus (P) in the litter and used denaturing gradient gel electrophoresis to investigate changes in fungal community composition. The C content of S. auriculatum litter did not change throughout the incubation period whereas vascular plant litters lost 30-40% of their initial C. There were no differences in C losses between low and medium water tables, but losses were always significantly less at the high water table. Most litters accumulated N and E. angustifolium accumulated significant quantities of P. C, N and P were significant explanatory variables in determining changes in fungal community composition but explained <25% of the variation. Litter type was always a stronger factor than water table in determining either fungal community composition or turnover of C, N and P in litter. The results have implications for the ways restoration programmes and global climate change may impact upon nutrient cycling in cutover peatlands.