Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically-limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net Nmin), i.e., the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net Nmin across a wide range of grasslands that vary in soil and climatic properties. Here, we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory-based) and realized (field-based) soil net Nmin. Herbivore removal in the absence of fertilization did not alter potential and realized soil net Nmin. However, fertilization alone and in combination with herbivore removal consistently increased potential soil net Nmin. Realized soil net Nmin, in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net Nmin were contingent on site-specific soil and climatic properties. Fertilization effects on potential soil net Nmin were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net Nmin declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion, and alterations in future climatic conditions can negatively impact soil net Nmin across global grasslands under realistic field conditions. This is important context-dependent knowledge for grassland management worldwide.
more »
« less
Direct and indirect effects of nitrogen enrichment on soil organisms and carbon and nitrogen mineralization in a semi‐arid grassland
Semi‐arid grasslands on the Mongolian Plateau are expected to experience high inputs of anthropogenic reactive nitrogen in this century. It remains unclear, however, how soil organisms and nutrient cycling are directly affected by N enrichment (i.e., without mediation by plant input to soil) vs. indirectly affected via changes in plant‐related inputs to soils resulting from N enrichment. To test the direct and indirect effects of N enrichment on soil organisms (bacteria, fungi and nematodes) and their associated C and N mineralization, in 2010, we designated two subplots (with plants and without plants) in every plot of a six‐level N‐enrichment experiment established in 1999 in a semi‐arid grassland. In 2014, 4 years after subplots with and without plant were established, N enrichment had substantially altered the soil bacterial, fungal and nematode community structures due to declines in biomass or abundance whether plants had been removed or not. N enrichment also reduced the diversity of these groups (except for fungi) and the soil C mineralization rate and induced a hump‐shaped response of soil N mineralization. As expected, plant removal decreased the biomass or abundance of soil organisms and C and N mineralization rates due to declines in soil substrates or food resources. Analyses of plant‐removal‐induced changes (ratios of without‐ to with‐plant subplots) showed that micro‐organisms and C and N mineralization rates were not enhanced as N enrichment increased but that nematodes were enhanced as N enrichment increased, indicating that the effects of plant removal on soil organisms and mineralization depended on trophic level and nutrient status. Surprisingly, there was no statistical interaction between N enrichment and plant removal for most variables, indicating that plant‐related inputs did not qualitatively change the effects of N enrichment on soil organisms or mineralization. Structural equation modelling confirmed that changes in soil communities and mineralization rates were more affected by the direct effects of N enrichment (via soil acidification and increased N availability) than by plant‐related indirect effects. Our results provide insight into how future changes in N deposition and vegetation may modify below‐ground communities and processes in grassland ecosystems.
more »
« less
- Award ID(s):
- 1557417
- PAR ID:
- 10094378
- Date Published:
- Journal Name:
- Functional ecology
- ISSN:
- 1365-2435
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
-
Abstract Soil extracellular enzymes are produced and excreted by soil microbial organisms. They catalyze many of the biochemical reactions that support crucial ecosystem functions like decomposition, organic matter mineralization, nutrient cycling, and C sequestration. Microbial communities that produce these enzymes are shaped in part by plant diversity and composition, but the relationships between plant communities and enzyme activities are not well resolved. Diverse plant communities may provide a diversity of soil organic matter inputs that support high microbial diversity and function. This would predict positive relationships between enzyme activities and plant diversity, particularly plant functional trait diversity that more closely reflects the diversity of soil inputs. Alternatively, dominant plant functional groups may shape microbial communities and activities, such as legumes that add N‐rich resources to soil. These impacts might be seen in microbes by the relative acquisition of different resources, such as the activity ratio of C‐acquiring β‐glucosidase and N‐acquiring N‐acetyl‐β‐D‐glucosaminidase. We use soil enzyme and plant community data from four separate grassland studies—two experiments and two observational community studies—to ask how plant diversity and functional composition affect enzyme activities. There were relationships between enzyme activities and plants for the observational community studies, but not the experimental studies, so plant–soil enzyme relationships may take time to emerge. At one community site, activities for four hydrolytic enzymes declined with plant Shannon diversity, and β‐glucosidase activity increased with greater functional trait diversity. The ratio of C‐acquiring β‐glucosidase and N‐acquiring N‐acetyl‐β‐D‐glucosaminidase activities increased with plant diversity and cover of nitrogen‐fixers, or declined with graminoid cover, suggesting that microbes reduced their relative allocation of resources to N acquisition when plant inputs made N more accessible. If lower activities of enzymes targeting C‐rich compounds are indicators of higher potential for C sequestration, then grassland plant communities with high taxonomic diversity may promote C storage. This could provide strong justification for management and restoration strategies that sustain high plant biodiversity. However, the inconsistent results, with enzyme activity unrelated to plants in both experimental studies, imply that other environmental factors correlated with plant composition may be stronger determinants of soil enzymes in some grassland settings.more » « less
-
Nitrogen (N) deposition increases soil carbon (C) storage by reducing microbial activity. These effects vary in soil beneath trees that associate with arbuscular (AM) and ectomycorrhizal (ECM) fungi. Variation in carbon C and N uptake traits among microbes may explain differences in soil nutrient cycling between mycorrhizal associations in response to high N loads, a mechanism not previously examined due to methodological limitations. Here, we used quantitative Stable Isotope Probing (qSIP) to measure bacterial C and N assimilation rates from an added organic compound, which we conceptualize as functional traits. As such, we applied a trait‐based approach to explore whether variation in assimilation rates of bacterial taxa can inform shifts in soil function under chronic N deposition. We show taxon‐specific and community‐wide declines of bacterial C and N uptake under chronic N deposition in both AM and ECM soils. N deposition‐induced reductions in microbial activity were mirrored by declines in soil organic matter mineralization rates in AM but not ECM soils. Our findings suggest C and N uptake traits of bacterial communities can predict C cycling feedbacks to N deposition in AM soils, but additional data, for instance on the traits of fungi, may be needed to connect microbial traits with soil C and N cycling in ECM systems. Our study also highlights the potential of employing qSIP in conjunction with trait‐based analytical approaches to inform how ecological processes of microbial communities influence soil functioning.more » « less
-
null (Ed.)which limit cell wall digestibility and efficiency of cellulose conversion to bioethanol, can be influenced by belowground biotic and abiotic factors. Switchgrass (Panicum virgatum L.) is a leading lignocellulosic biofuel crop and forms strong belowground associations with arbuscular mycorrhizal fungi (AMF), is susceptible to belowground plant-parasitic nematodes (PPN), and when grown in monoculture generally requires nitrogen (N) fertilization. The main objectives of the study were to investigate the effects of N fertilizer and belowground organisms on lignin content and composition of switchgrass. Leaf, stem, and root tissues were evaluated separately to test whether these factors had varying belowground (local) or aboveground (systemic) effects on plants. These factors were manipulated in a field study in 2017 using biocide applications to reduce soil fungi and nematodes. Combined biocide application reduced p-hydroxyphenyl (H) unit abundance in the leaves by 14% and increased the syringyl:guaiacyl (S:G) ratio in stems by 2%. Application of fungicide alone increased stem syringyl (S) unit by 12.4% as compared with control plots, and 11.1% as compared with nematicide plots. Overall, fertilizer increased total stem lignin by 3%, stem S unit by 6.7%, and stem S:G ratio by 10%, whereas it reduced the amount of H-unit in the roots by 11%. While the effects of N fertilizer were more pronounced in this study, changes to soil organisms had similar magnitudes of effect for some measures of lignin, indicating that these belowground interactions may be important for growers to consider in the future.more » « less
-
null (Ed.)Whether the terrestrial biosphere will continue to act as a net carbon (C) sink in the face of multiple global changes is questionable. A key uncertainty is whether increases in plant C fixation under elevated carbon dioxide (CO 2 ) will translate into decades-long C storage and whether this depends on other concurrently changing factors. We investigated how manipulations of CO 2 , soil nitrogen (N) supply, and plant species richness influenced total ecosystem (plant + soil to 60 cm) C storage over 19 y in a free-air CO 2 enrichment grassland experiment (BioCON) in Minnesota. On average, after 19 y of treatments, increasing species richness from 1 to 4, 9, or 16 enhanced total ecosystem C storage by 22 to 32%, whereas N addition of 4 g N m −2 ⋅ y −1 and elevated CO 2 of +180 ppm had only modest effects (increasing C stores by less than 5%). While all treatments increased net primary productivity, only increasing species richness enhanced net primary productivity sufficiently to more than offset enhanced C losses and substantially increase ecosystem C pools. Effects of the three global change treatments were generally additive, and we did not observe any interactions between CO 2 and N. Overall, our results call into question whether elevated CO 2 will increase the soil C sink in grassland ecosystems, helping to slow climate change, and suggest that losses of biodiversity may influence C storage as much as or more than increasing CO 2 or high rates of N deposition in perennial grassland systems.more » « less
An official website of the United States government

