Does the 2013 GOLD classification improve the ability to predict lung function decline, exacerbations and mortality: a post-hoc analysis of the 4-year UPLIFT trial

BACKGROUND: The 2013 GOLD classification system for COPD distinguishes four stages: A (low symptoms, low exacerbation risk), B (high symptoms, low risk), C (low symptoms, high risk) and D (high symptoms, high risk). Assessment of risk is based on exacerbation history and airflow obstruction, whatever results in a higher risk grouping. The previous system was solely based on airflow obstruction. Earlier studies compared the predictive performance of new and old classification systems with regards to mortality and exacerbations. The objective of this study was to compare the ability of both classifications to predict the number of future (total and severe) exacerbations and mortality in a different patient population, and to add an outcome measure to the comparison: lung function decline.METHODS: Patient-level data from the UPLIFT trial were used to analyze 4-year survival in a Weibull model, with GOLD stages at baseline as covariates. A generalized linear model was used to compare the numbers of exacerbations (total and severe) per stage. Analyses were repeated with stages C and D divided into substages depending on lung function and exacerbation history. Lung function decline was analysed in a repeated measures model.RESULTS: Mortality increased from A to D, but there was no difference between B and C. For the previous GOLD stages 2-4, survival curves were clearly separated. Yearly exacerbation rates were: 0.53, 0.72 and 0.80 for stages 2-4; and 0.35, 0.45, 0.58 and 0.74 for A-D. Annual rates of lung function decline were: 47, 38 and 26 ml for stages 2-4 and 44, 48, 38 and 39 for stages A-D. With regards to model fit, the new system performed worse at predicting mortality and lung function decline, and better at predicting exacerbations. Distinguishing between the sub-stages of high-risk led to substantial improvements.CONCLUSIONS: The new classification system is a modest step towards a phenotype approach. It is probably an improvement for the prediction of exacerbations, but a deterioration for predicting mortality and lung function decline.TRIAL REGISTRATION: ClinicalTrials.gov NCT00144339 (September 2, 2005)

Land‐use intensity and biodiversity effects on infiltration capacity and hydraulic conductivity of grassland soils in southern Germany

Evidence from experimental and established grasslands indicates that plant biodiversity can modify the water cycle. One suspected mechanism behind this is a higher infiltration capacity (ν$_{B}$) and hydraulic conductivity (K) of the soil on species-rich grasslands. However, in established and agriculturally managed grasslands, biodiversity effects cannot be studied independent of land-use effects. Therefore, we investigated in established grassland systems how land-use intensity and associated biodiversity of plants and soil animals affect νB and K at and close to saturation. On 50 grassland plots along a land-use intensity gradient in the Biodiversity Exploratory Schwäbische Alb, Germany, we measured νB with a hood infiltrometer at several matrix potentials and calculated the saturated and unsaturated K. We statistically analysed the relationship between ν$_{B}$ or K and land-use information (e.g., fertilising intensity), abiotic (e.g., soil texture) and biotic data (e.g., plant species richness, earthworm abundance). Land-use intensity decreased and plant species richness increased ν$_{B}$ and K, while the direction of the effects of soil animals was inconsistent. The effect of land-use intensity on ν$_{B}$ and K was mainly attributable to its negative effect on plant species richness. Our results demonstrate that plant species richness was a better predictor of ν$_{B}$ and K at and close to saturation than land-use intensity or soil physical properties in the established grassland systems of the Schwäbische Alb

Biodiversity effects on ecosystem functioning in a 15-year grassland experiment: Patterns, mechanisms, and open questions

In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research. First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the ‘main experiment’, where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity–stability theory. Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness. Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above- than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances. Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, element turnover, element stocks, and output from the ecosystem. While inputs were generally less affected by plant species richness, measures of element stocks, turnover and output were often positively affected by plant diversity, e.g. carbon storage strongly increased with increasing plant species richness. Variables of the N cycle responded less strongly to plant species richness than variables of the C cycle. Fifth, plant traits are often used to unravel mechanisms underlying the biodiversity–ecosystem functioning relationship. In the Jena Experiment, most investigated plant traits, both above- and belowground, were plastic and trait expression depended on plant diversity in a complex way, suggesting limitation to using database traits for linking plant traits to particular functions. Sixth, plant diversity effects on ecosystem processes are often caused by plant diversity effects on species interactions. Analyses in the Jena Experiment including structural equation modelling suggest complex interactions that changed with diversity, e.g. soil carbon storage and greenhouse gas emission were affected by changes in the composition and activity of the belowground microbial community. Manipulation experiments, in which particular organisms, e.g. belowground invertebrates, were excluded from plots in split-plot experiments, supported the important role of the biotic component for element and water fluxes. Seventh, the Jena Experiment aimed to put the results into the context of agricultural practices in managed grasslands. The effect of increasing plant species richness from 1 to 16 species on plant biomass was, in absolute terms, as strong as the effect of a more intensive grassland management, using fertiliser and increasing mowing frequency. Potential bioenergy production from high-diversity plots was similar to that of conventionally used energy crops. These results suggest that diverse ‘High Nature Value Grasslands’ are multifunctional and can deliver a range of ecosystem services including production-related services. A final task was to assess the importance of potential artefacts in biodiversity–ecosystem functioning relationships, caused by the weeding of the plant community to maintain plant species composition. While the effort (in hours) needed to weed a plot was often negatively related to plant species richness, species richness still affected the majority of ecosystem variables. Weeding also did not negatively affect monoculture performance; rather, monocultures deteriorated over time for a number of biological reasons, as shown in plant-soil feedback experiments. To summarize, the Jena Experiment has allowed for a comprehensive analysis of the functional role of biodiversity in an ecosystem. A main challenge for future biodiversity research is to increase our mechanistic understanding of why the magnitude of biodiversity effects differs among processes and contexts. It is likely that there will be no simple answer. For example, among the multitude of mechanisms suggested to underlie the positive plant species richness effect on biomass, some have received limited support in the Jena Experiment, such as vertical root niche partitioning. However, others could not be rejected in targeted analyses. Thus, from the current results in the Jena Experiment, it seems likely that the positive biodiversity effect results from several mechanisms acting simultaneously in more diverse communities, such as reduced pathogen attack, the presence of more plant growth promoting organisms, less seed limitation, and increased trait differences leading to complementarity in resource uptake. Distinguishing between different mechanisms requires careful testing of competing hypotheses. Biodiversity research has matured such that predictive approaches testing particular mechanisms are now possible

Mechanisms behind plant diversity effects on inorganic and organic N leaching from temperate grassland

Higher plant diversity reduces nitrate leaching by complementary resource use, while its relation to leaching of other N species is unclear. We determined the effects of plant species richness, functional group richness, and the presence of specific functional groups on ammonium, dissolved organic N (DON), and total dissolved N (TDN) leaching from grassland in the first 4 years after conversion from fertilized arable land to unfertilized grassland. On 62 experimental plots in Jena, Germany, with 1–60 plant species and 1–4 functional groups (legumes, grasses, tall herbs, small herbs), nitrate, ammonium, and TDN concentrations in soil solution (0–0.3 m soil layer) were measured fortnightly during 4 years. DON concentrations were calculated by subtracting inorganic N from TDN. Nitrogen concentrations were multiplied with modeled downward water fluxes to obtain N leaching. DON leaching contributed most to TDN leaching (64 ± SD 4% of TDN). Ammonium leaching was unaffected by plant diversity. Increasing species richness decreased DON leaching in the fourth year. We attribute this finding to enhanced use of DON as a C and N source and enhanced mineralization of DON by soil microorganisms. An increase of species richness decreased TDN leaching likely driven by the complementary use of nitrate by diverse mixtures. Legumes increased DON and TDN leaching likely because of their N2-fixing ability and higher litter production. Grasses decreased TDN leaching because of more exhaustive use of nitrate and water. Our results demonstrate that increasing plant species richness decreases leaching of DON and TDN

Plant diversity enhances the natural attenuation of polycyclic aromatic compounds (PAHs and oxygenated PAHs) in grassland soils

Increasing plant species richness stimulates microbial activity in soil, which might favor biodegradation of polycyclic aromatic compounds (PACs). To explore the relationship between plant community composition and PACs in grassland soils (Fluvisols exposed to an urban atmosphere), we determined the concentrations of 29 polycyclic aromatic hydrocarbons (PAHs) and 15 oxygenated PAHs (OPAHs) in topsoils of 80 plots of a grassland biodiversity experiment. The plots included different levels of plant species richness (1, 2, 4, 8, 16, 60 species) and 1–4 plant functional groups (grasses, small herbs, tall herbs, and legumes) in a randomized block design. The concentrations (ng g−1) of ∑29PAHs and ∑15OPAHs in the soils were 271–2407 and 57–329, respectively. Concentrations of 16 (out of 44) PACs and ∑29PAHs decreased significantly with increasing plant species richness, after accounting for the effects of block and initial soil organic C concentration (ANCOVA, p 4 aromatic rings on plant species richness yielded slopes that were negatively correlated with their octanol-water partitioning coefficients, (ii) two OPAHs accumulated in soils with higher plant species richness, and (iii) higher plant species richness increased four OPAH/parent-PAH ratios. Accordingly, structural equation modeling indicated that the higher concentration of 1,2-acenaphthenequinone (a metabolite of acenaphthene) and the higher 1,2-acenaphthenequinone/acenaphthene and 1-indanone/fluorene ratios in plots with higher plant species richness were partly explained by higher soil microbial biomass on plots with higher plant species richness. We conclude that higher plant species richness can be used to enhance biodegradation of aged PACs in soil. We however caution that OPAHs (some of which are more toxic than their related PAHs) might accumulate in soils during such a plant-assisted remediation process