Effects of extreme rainfall events are independent of plant species richness in an experimental grassland community.

Global climate models predict more frequent periods of drought stress alternated by heavier, but fewer rainfall events in the future. Biodiversity studies have shown that such changed drought stress may be mitigated by plant species richness. Here, we investigate if grassland communities, differing in species richness, respond differently to climatic extremes within the growing season. In a 3-year outdoor mesocosm experiment, four grassland species in both monoculture and mixture were subjected to a rainfall distribution regime with two levels: periods of severe drought in the summer intermitted by extreme rainfall events versus regular rainfall over time. Both treatments received the same amount of water over the season. Extreme rainfall combined with drought periods resulted in a 15% decrease in aboveground biomass in the second and third year, compared to the regular rainfall regime. Root biomass was also reduced in the extreme rainfall treatment, particularly in the top soil layer (- 40%). All species developed higher water use efficiencies (less negative leaf δ13C) in extreme rainfall than in regular rainfall. These responses to the rainfall/drought treatment were independent of species richness, although the mixtures were on an average more productive in terms of biomass than the monocultures. Our experimental results suggest that mixtures are similarly able to buffer these within-season rainfall extremes than monocultures, which contrasts with findings in the studies on natural droughts. Our work demonstrates the importance of investigating the interactions between rainfall distribution and drought periods for understanding effects of climate change on plant community performance.

Spatial heterogeneity of plant-soil feedback affects root interactions and interspecific competition.

Plant-soil feedback is receiving increasing interest as a factor influencing plant competition and species coexistence in grasslands. However, we do not know how spatial distribution of plant-soil feedback affects plant below-ground interactions. We investigated the way in which spatial heterogeneity of soil biota affects competitive interactions in grassland plant species. We performed a pairwise competition experiment combined with heterogeneous distribution of soil biota using four grassland plant species and their soil biota. Patches were applied as quadrants of 'own' and 'foreign' soils from all plant species in all pairwise combinations. To evaluate interspecific root responses, species-specific root biomass was quantified using real-time PCR. All plant species suffered negative soil feedback, but strength was species-specific, reflected by a decrease in root growth in own compared with foreign soil. Reduction in root growth in own patches by the superior plant competitor provided opportunities for inferior competitors to increase root biomass in these patches. These patterns did not cascade into above-ground effects during our experiment. We show that root distributions can be determined by spatial heterogeneity of soil biota, affecting plant below-ground competitive interactions. Thus, spatial heterogeneity of soil biota may contribute to plant species coexistence in species-rich grasslands.

Virus infection decreases the attractiveness of white clover plants for a non-vectoring herbivore.

Plant pathogens and insect herbivores are prone to share hosts under natural conditions. Consequently, pathogen-induced changes in the host plant can affect herbivory, and vice versa. Even though plant viruses are ubiquitous in the field, little is known about plant-mediated interactions between viruses and non-vectoring herbivores. We investigated the effects of virus infection on subsequent infestation by a non-vectoring herbivore in a natural genotype of Trifolium repens (white clover). We tested whether infection with White clover mosaic virus (WClMV) alters (1) the effects of fungus gnat feeding on plant growth, (2) the attractiveness of white clover for adult fungus gnat females, and (3) the volatile emission of white clover plants. We observed only marginal effects of WClMV infection on the interaction between fungus gnat larvae and white clover. However, adult fungus gnat females clearly preferred non-infected over WClMV-infected plants. Non-infected and virus-infected plants could easily be discriminated based on their volatile blends, suggesting that the preference of fungus gnats for non-infected plants may be mediated by virus-induced changes in volatile emissions. The compound ß-caryophyllene was exclusively detected in the headspace of virus-infected plants and may hence be particularly important for the preference of fungus gnat females. Our results demonstrate that WClMV infection can decrease the attractiveness of white clover plants for fungus gnat females. This suggests that virus infections may contribute to protecting their hosts by decreasing herbivore infestation rates. Consequently, it is conceivable that viruses play a more beneficial role in plant-herbivore interactions than generally thought.

Interspecific and intraspecific differences in shoot and leaf lifespan of four Carex species which differ in maximum dry matter production.

The effect of N supply on shoot and leaf lifespan was investigated in established stands of four herbaceous Carex species which differed in maximum dry matter production. These species were, in rank order of increasing maximum dry matter production (per unit ground area): Carex diandra≤C. rostrata

Nitrous oxide emission and denitrification in chronically nitrate-loaded riparian buffer zones.

Riparian buffer zones are known to reduce diffuse N pollution of streams by removing and modifying N from agricultural runoff. Denitrification, often identified as the key N removal process, is also considered as a major source of the greenhouse gas nitrous oxide (N2O). The risks of high N2O emissions during nitrate mitigation and the environmental controls of emissions have been examined in relatively few riparian zones and the interactions between controls and emissions are still poorly understood. Our objectives were to assess the rates of N2O emission from riparian buffer zones that receive large loads of nitrate, and to evaluate various factors that are purported to control N emissions. Denitrification, nitrification, and N2O emissions were measured seasonally in grassland and forested buffer zones along first-order streams in The Netherlands. Lateral nitrate loading rates were high, up to 470 g N m(-2) yr(-1). Nitrogen process rates were determined using flux chamber measurements and incubation experiments. Nitrous oxide emissions were found to be significantly higher in the forested (20 kg N ha(-1) yr(-1)) compared with the grassland buffer zone (2-4 kg N ha(-1) yr(-1)), whereas denitrification rates were not significantly different. Higher rates of N2O emissions in the forested buffer zone were associated with higher nitrate concentrations in the ground water. We conclude that N transformation by nitrate-loaded buffer zones results in a significant increase of greenhouse gas emission. Considerable N2O fluxes measured in this study indicate that Intergovernmental Panel on Climate Change methodologies for quantifying indirect N2O emissions have to distinguish between agricultural uplands and riparian buffer zones in landscapes receiving large N inputs.

Early root overproduction not triggered by nutrients decisive for competitive success belowground.

BACKGROUND: Theory predicts that plant species win competition for a shared resource by more quickly preempting the resource in hotspots and by depleting resource levels to lower concentrations than its competitors. Competition in natural grasslands largely occurs belowground, but information regarding root interactions is limited, as molecular methods quantifying species abundance belowground have only recently become available. PRINCIPAL FINDINGS: In monoculture, the grass Festuca rubra had higher root densities and a faster rate of soil nitrate depletion than Plantago lanceolata, projecting the first as a better competitor for nutrients. However, Festuca lost in competition with Plantago. Plantago not only replaced the lower root mass of its competitor, but strongly overproduced roots: with only half of the plants in mixture than in monoculture, Plantago root densities in mixture were similar or higher than those in its monocultures. These responses occurred equally in a nutrient-rich and nutrient-poor soil layer, and commenced immediately at the start of the experiment when root densities were still low and soil nutrient concentrations high. CONCLUSIONS/SIGNIFICANCE: Our results suggest that species may achieve competitive superiority for nutrients by root growth stimulation prior to nutrient depletion, induced by the presence of a competitor species, rather than by a better ability to compete for nutrients per se. The root overproduction by which interspecific neighbors are suppressed independent of nutrient acquisition is consistent with predictions from game theory. Our results emphasize that root competition may be driven by other mechanisms than is currently assumed. The long-term consequences of these mechanisms for community dynamics are discussed.

Effects of cell number and cell size on petiole length variation in a stoloniferous herb.

In stoloniferous species, the length of petioles is of pivotal importance because it determines the position of leaf blades within the canopy. From a mechanistic perspective, two developmental processes, cell division and cell elongation, are responsible for the length of a given petiole. This study aimed at quantifying the relative contributions of cell division and cell elongation to genotypic and plastic variation in petiole length of the stoloniferous herb Trifolium repens. Thirty-four genotypes of T. repens were grown under high light conditions and simulated canopy shade. Cells were counted and their lengths measured on epidermal prints from fully grown petioles of leaves that had been initiated in the experimental light conditions. Cell number was the main trait explaining petiole length differences among genotypes grown under high light, while both cell number and length changed in response to shading. Our study revealed a strong negative correlation between shade-induced changes in cell number and cell length: genotypes that responded to shading by increasing cell numbers hardly changed in cell length, and vice versa. Our results suggest that genotypic and phenotypic variation in petiole length results from a complex interplay between the developmental processes of cell elongation and cell division.