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Abstract Grasslands cover approximately a third of the Earth’s land surface and account for about a third of terrestrial carbon storage. Yet, we lack strong predictive models of grassland plant biomass, the primary source of carbon in grasslands. This lack of predictive ability may arise from the assumption of linear relationships between plant biomass and the environment and an underestimation of interactions of environmental variables. Using data from 116 grasslands on six continents, we show unimodal relationships between plant biomass and ecosystem characteristics, such as mean annual precipitation and soil nitrogen. Further, we found that soil nitrogen and plant diversity interacted in their relationships with plant biomass, such that plant diversity and biomass were positively related at low levels of nitrogen and negatively at elevated levels of nitrogen. Our results show that it is critical to account for the interactive and unimodal relationships between plant biomass and several environmental variables to accurately include plant biomass in global vegetation and carbon models. 

Abstract Plant disease often increases with N, decreases with CO₂, and increases as biodiversity is lost (i.e., the dilution effect). Additionally, all these factors can indirectly alter disease by changing host biomass and hence density-dependent disease transmission. Yet over long periods of time as communities undergo compositional changes, these biomass-mediated pathways might fade, intensify, or even reverse in direction. Using a field experiment that has manipulated N, CO₂, and species richness for over 20 years, we compared severity of a specialist rust fungus (Puccinia andropogonis) on its grass host (Andropogon gerardii) shortly after the experiment began (1999) and twenty years later (2019). Between these two sampling periods, two decades apart, we found that disease severity consistently increased with N and decreased with CO₂. However, the relationship between diversity and disease reversed from a dilution effect in 1999 (more severe disease in monocultures) to an amplification effect in 2019 (more severe disease in mixtures). The best explanation for this reversal centered on host density (i.e., aboveground biomass), which was initially highest in monoculture, but became highest in mixtures two decades later. Thus, the diversity-disease pattern reversed, but disease consistently increased with host biomass. These results highlight the consistency of N and CO₂as drivers of plant disease in the Anthropocene and emphasize the critical role of host biomass—despite potentially variable effects of diversity—for relationships between biodiversity and disease. 

Plants serve as critical links between above- and below-ground microbial communitites, both influencing and being influenced by microbes in these two realms. Below-ground microbial communities are expected to respond to soil resource environments, which are mediated by the roots of plants that can, in turn, be influenced by the above-ground community of foliar endophytes. For instance, diverse plant communities deposit more, and more diverse, nutrients into the soil, and this deposition is often increased when foliar pathogens are removed. Differences in soil resources can alter soil microbial composition and phenotypes, including inhibitory capacity, resource use, and antibiotic resistance. In this work, we consider plots differing in plant richness and application of foliar fungicide, evaluating consequences on soil resource levels and root-associatedStreptomycesphenotypes. Soil carbon, nitrogen, phosphorus, potassium, and organic matter were greater in samples from polyculture than monoculture, yet this increase was surprisingly offset when foliar fungal communities were disrupted. We find thatStreptomycesphenotypes varied more between richness plots—with theStreptomycesfrom polyculture showing lower inhibitory capacity, altered resource-use profiles, and greater antibiotic resistance—than between subplots with/without foliar fungicide. Where foliar fungicide affected phenotypes, it did so differently in polyculture than in monoculture, for instance decreasing niche width and overlap in monoculture while increasing them in polyculture. No differences in phenotype were correlated with soil nutrient levels, suggesting the need for further research looking more closely at soil resource diversity and particular compounds that were found to differ between treatments. 

Experiments comparing diploids with polyploids and in single grassland sites show that nitrogen and/or phosphorus availability influences plant growth and community composition dependent on genome size; specifically, plants with larger genomes grow faster under nutrient enrichments relative to those with smaller genomes. However, it is unknown if these effects are specific to particular site localities with speciifc plant assemblages, climates, and historical contingencies. To determine the generality of genome size-dependent growth responses to nitrogen and phosphorus fertilization, we combined genome size and species abundance data from 27 coordinated grassland nutrient addition experiments in the Nutrient Network that occur in the Northern Hemisphere across a range of climates and grassland communities. We found that after nitrogen treatment, species with larger genomes generally increased more in cover compared to those with smaller genomes, potentially due to a release from nutrient limitation. Responses were strongest for C₃grasses and in less seasonal, low precipitation environments, indicating that genome size effects on water-use-efficiency modulates genome size–nutrient interactions. Cumulatively, the data suggest that genome size is informative and improves predictions of species’ success in grassland communities. 

Abstract A combination of theory and experiments predicts that increasing soil nutrients will modify herbivore and microbial impacts on ecosystem carbon cycling.However, few studies of herbivores and soil nutrients have measured both ecosystem carbon fluxes and carbon pools. Even more rare are studies manipulating microbes and nutrients that look at ecosystem carbon cycling responses.We added nutrients to a long‐term, experiment manipulating foliar fungi, soil fungi, mammalian herbivores and arthropods in a low fertility grassland. We measured gross primary production (GPP), ecosystem respiration (ER), net ecosystem exchange (NEE) and plant biomass throughout the growing season to determine how nutrients modify consumer impacts on ecosystem carbon cycling.Nutrient addition increased above‐ground biomass and GPP, but not ER, resulting in an increase in ecosystem carbon uptake rate. Reducing foliar fungi and arthropods increased plant biomass. Nutrients amplified consumer effects on plant biomass, such that arthropods and foliar fungi had a threefold larger impact on above‐ground biomass in fertilized plots.Synthesis. Our work demonstrates that throughout the growing season soil resources modify carbon uptake rates as well as animal and fungal impacts on plant biomass production. Taken together, ongoing nutrient pollution may increase ecosystem carbon uptake and drive fungi and herbivores to have larger impacts on plant biomass production. 

Eukaryotic hosts harbor tremendously diverse microbiomes that affect host fitness and response to environmental challenges. Fungal endophytes are prominent members of plant microbiomes, but we lack information on the diversity in functional traits affecting their interactions with their host and environment. We used two culturing approaches to isolate fungal endophytes associated with the widespread, dominant prairie grassAndropogon gerardiiand characterized their taxonomic diversity using rDNA barcode sequencing. A randomly chosen subset of fungi representing the diversity of each leaf was then evaluated for their use of different carbon compound resources and growth on those resources. Applying community phylogenetic analyses, we discovered that these fungal endophyte communities are comprised of phylogenetically distinct assemblages of slow- and fast-growing fungi that differ in their use and growth on differing carbon substrates. Our results demonstrate previously undescribed and cryptic functional diversity in carbon resource use and growth in fungal endophyte communities ofA.gerardii. 

Death is a common outcome of infection, but most disease models do not track hosts after death. Instead, these hosts disappear into a void. This assumption lacks critical realism, because dead hosts can alter host–pathogen dynamics. Here, we develop a theoretical framework of carbon‐based models combining disease and ecosystem perspectives to investigate the consequences of feedbacks between living and dead hosts on disease dynamics and carbon cycling. Because autotrophs (i.e. plants and phytoplankton) are critical regulators of carbon cycling, we developed general model structures and parameter combinations to broadly reflect disease of autotrophic hosts across ecosystems. Analytical model solutions highlight the importance of disease–ecosystem coupling. For example, decomposition rates of dead hosts mediate pathogen spread, and carbon flux between live and dead biomass pools are sensitive to pathogen effects on host growth and death rates. Variation in dynamics arising from biologically realistic parameter combinations largely fell along a single gradient from slow to fast carbon turnover rates, and models predicted higher disease impacts in fast turnover systems (e.g. lakes and oceans) than slow turnover systems (e.g. boreal forests). Our results demonstrate that a unified framework, including the effects of pathogens on carbon cycling, provides novel hypotheses and insights at the nexus of disease and ecosystem ecology. 

Abstract Plants face trade‐offs between allocating resources to growth, while also defending against herbivores or pathogens. Species differences along defense trade‐off axes may promote coexistence and maintain diversity. However, few studies of plant communities have simultaneously compared defense trade‐offs against an array of herbivores and pathogens for which defense investment may differ, and even fewer have been conducted in the complex natural communities in which these interactions unfold. We tested predictions about the role of defense trade‐offs with competition and growth in diversity maintenance by tracking plant species abundance in a field experiment that removed individual consumer groups (mammals, arthropods, fungi) and added nutrients. Consistent with a growth–defense trade‐off, plant species that increased in mass in response to nutrient addition also increased when consumers were removed. This growth–defense trade‐off occurred for all consumer groups studied. Nutrient addition reduced plant species richness, which is consistent with trade‐off theory. Removing foliar fungi increased plant diversity via increased species evenness, whereas removal of other consumer groups had little effect on diversity, counter to expectations. Thus, while growth–defense trade‐offs are general across consumer groups, this trade‐off observed in wild plant communities does not necessarily support plant diversity maintenance. 

Abstract Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance.