Anthropogenic effects on global soil nitrogen pools.

The amount of nitrogen stored in terrestrial soils, its "nitrogen pool", moderates biogeochemical cycling affecting primary productivity, nitrogen pollution and even carbon budgets. The soil nitrogen pools and the transformation of nitrogen forms within them are heavily influenced by environmental factors including anthropogenic activities. However, our understanding of the global distribution of soil nitrogen with respect to anthropogenic activity and human land use remains unclear. We constructed a meta-analysis from a global sampling, in which we compare soil total nitrogen pools and the driving mechanisms affecting each pool across three major classifications of human land use: natural, agricultural, and urban. Although the size of the nitrogen pool can be similar across natural, agricultural and urban soils, the ecological and human associated drivers vary. Specifically, the drivers within agricultural and urban soils as opposed to natural soils are more complex and often decoupled from climatic and soil factors. This suggests that the nitrogen pools of those soils may be co-moderated by other factors not included in our analyses, like human activities. Our analysis supports the notion that agricultural soils act as a nitrogen source while urban soils as a nitrogen sink and informs a modern understanding of the fates and distributions of anthropogenic nitrogen in natural, agricultural, and urban soils.

Mycorrhizal Infection Can Ameliorate Abiotic Factors in Urban Soils.

Once abandoned, urban and post-industrial lands can undergo a re-greening, the natural regeneration and succession that leads to surprisingly healthy plant communities, but this process is dependent upon microbial activity and the health of the parent soil. This study aimed to evaluate the effects of arbuscular mycorrhizal fungi (AMF) in facilitating plant production in post-industrial soils. In so doing, we helped to resolve the mechanism through which AMF ameliorate environmental stress in terrestrial plants. An experiment was established in which rye grass (Lolium perenne) was grown in two heavy metal-contaminated soils from an urban brownfield in New Jersey, USA, and one non-contaminated control soil. One set of the treatments received an AMF inoculum (four species in a commercial mix: Glomus intraradices, G. mosseae, G. etunicatum and G. aggregatum) and the other did not. Upon harvest, dried plant biomass, root/shoot ratio, AMF colonization, and extracellular soil phosphatase activity, a proxy for soil microbial functioning, were all measured. Plant biomass increased across all treatments inoculated with AMF, with a significantly higher average shoot and root mass compared to non-inoculated treatments. AMF colonization of the roots in contaminated soil was significantly higher than colonization in control soil, and the root/shoot ratio of plants in contaminated soils was also higher when colonized by AMF. Mycorrhizal infection may help plants to overcome the production limits of post-industrial soils as is seen here with increased infection and growth. The application of this mechanistic understanding to remediation and restoration strategies will improve soil health and plant production in urban environments.

Artificial root exudates restore microbial functioning in a metal contaminated, barren, inactive soil.

Restoring enzyme function in barren, brownfield soils using green strategies can improve microbial functioning and enable phytoremediation. It is known that adding simple, readily metabolized substrates secreted by growing plant roots (root exudates) or a laboratory prepared solution of root exudates (artificial root exudates) can stimulate soil microbial function. It is not known whether and how well this strategy works in a contaminated, low functioning soil from an industrial barren site because contaminants in the barren soil might inhibit microbial survival and functioning, or the microbial community might not be adapted to functionally benefit from root exudates. The objective of this study was to determine whether artificial root exudates stimulate microbial function in a barren soil. We collected soils from a barren brownfield (25R) site and an adjacent vegetated brownfield site (25F), with low and high enzyme activities, respectively. We subjected both soils to three treatments: switchgrass (native to the site), artificial root exudates, and a combination of switchgrass and artificial root exudates. We measured enzymatic activity, plant growth, soil moisture, organic matter content, and easily extractable glomalin content over 205 days. By day 157, artificial root exudates increased the phosphatase activity by 9-fold in previously vegetated brownfield soil and by 351-fold in barren brownfield soil. When exudates were added to the barren soil, the plant shoot mass was higher (52.2 ± 2.5 mg) than when they were not (35.4 ± 3.6 mg). In both soils, adding artificial root exudates significantly increased the percent moisture, organic matter, and glomalin content. Treating contaminated, barren soil with artificial root exudates resulted in increased soil microbial function and improved soil properties that might promote a hospitable habitat to support vegetation in such extreme environments. Summary: We added artificial root exudates to stimulate enzymatic function in two contaminated soils. Plant shoot mass, soil percent moisture, glomalin content, and organic matter content significantly increased due to the addition of artificial root exudates to the study soils. Microbially-mediated phosphatase activity was established in a barren, previously inactive, polluted soil.

Natural versus urban global soil organic carbon stocks: A meta-analysis.

Increasingly, the human existence in urban environments is growing. In addition, anthropogenic activity has altered the global carbon (C) cycle and triggered climate change. Soil is the largest pool of organic C in terrestrial ecosystems, but its ability to retain and store C varies. As humans move forward to mitigate climate change, there is a growing need to understand the C storing capacity of soils and their interactions with factors like climate, vegetation or a footprint of human activity. Here, we constructed a meta-analysis which focused on 30 cm soil depth by collecting data from over 191 studies measuring soil organic carbon (SOC) stocks across natural, urban green space, and urban intensive habitats. We then compared the SOC data between different climatic zones, vegetation types, and anthropogenic influences with the human footprint index. The results indicate that SOC stocks in natural habitats (98.22 ± 49.10 Mg ha-1) are significantly higher than those of urban green spaces (54.61 ± 22.02 Mg ha-1) and urban intensive habitats (65.88 ± 35.27 Mg ha-1). We find a significant and negative relationship between the human footprint and SOC stocks of natural habitats but not between the human footprint and either of the urban habitats. Urban intensive and urban green space habitat soils store less C than natural ones. However, when compared across climatic zones or vegetation types, the capacity of natural soils to store C is variable and vulnerable to human activity. Carbon storage in urban soils is likely limited by persistent and stable anthropogenic influences keeping variability low. This is most pronounced in urban green spaces where human management is high (i.e. a golf course) and SOC is low. A comprehensive understanding of C storage in soils is essential to land management and climate mitigation measures.

Grazing influences biomass production and protein content of alpine meadows.

Alpine grasslands are essential for carbon sequestration and food supply for domestic and wild herbivores inhabiting mountainous areas worldwide. These biomes, however, are alternatively threatened by the abandonment of agricultural and livestock practices leading to a fast-growing shrubification process while other mountain grasslands are suffering from the impacts of overgrazing. The functioning of alpine meadow ecosystems is primarily driven by climatic conditions, land-use legacies and grazing. However, although it is critically important, the role of large herbivores on the aboveground biomass and protein content of palatable plants is poorly understood for most alpine meadows. In this work, we explore the effects of grazing on grassland vegetation at two different spatial and temporal scales in the Eastern Pyrenees, Spain. Remote sensing was used to assess the effect of high and moderate grazing (HG and MG respectively) on grass biomass using the leaf area index (LAI) at the meso-scale (patches between 2.3 and 38.7 ha). We also explored the impact of null (NG), overgrazing (MO, mimicked overgrazing) and high (HG) grazing intensities at local scale setting eighteen 1 m2 exclusion boxes in six meadows (three boxes each) commonly used by domestic and wild ungulates. Historical satellite data showed that LAI values are greater in high than in low grazed areas (HG, mean = 0.66, LG, mean = 0.55). Along the same lines, high and moderate grazing pressures improved biomass production at the local-scale (HG, mean = 590.3 g/m2, MO, mean = 389.3 g/m2 and NG, mean = 110.8 g/m2). Crude protein content reached higher values under MO pressure than under HG pressure. Our results confirm that grazing intensity exerts significant changes on the above-ground biomass production and the protein content of plants consumed by domestic (cattle and horses) and wild ungulates (Southern Chamois, Rupicapra pyrenaica). We can conclude that ungulates sustain biomass and nutritive values of grass exerting a negligible effect on biomass and protein content of woody vegetation. Our results will inform management guidelines to support profitable grazing activities and promote conservation of the open landscapes in the alpine ecosystems under the current global change scenario.

Plants mitigate restrictions to phosphatase activity in metal contaminated soils.

Soil anthropogenic contaminants can limit enzymatic nutrient mineralization, either by direct regulation or via impacts on the microbial community, thus affecting plant growth in agricultural and non-agricultural soils. The impact on phosphatase activity of mixing two contaminated, post-industrial rail yard soils was investigated; one was vegetated and had high phosphatase function, the other was barren and had low enzymatic function. The two soils had different abiotic properties, including contaminant load, vegetation cover, soil aggregate size distribution, and phosphatase potential. An experimental gradient was established between the two soils to systematically vary the abiotic properties and microbial community composition of the two soils, creating a gradient of novel ecosystems. The time dependence of extracellular phosphatase activity, soil moisture, and organic matter content was assessed along this gradient in the presence and absence of plants. Initially, mixtures with higher percentages of functional, vegetated soil had higher phosphatase activities. Phosphatase activity remained unchanged through time (65 days) in all soil mixtures in unplanted pots, but it increased in planted pots. For example, in the presence of plants, phosphatase activity increased from 0.6 ± 0.1 to 2.4 ± 0.3 µmol•h-1•gdry soil-1 from day one to day 65 in the 1:1 functional:barren soil mixture. The presence of plants also promoted moisture retention. Inoculation of poorly functioning soil with 10% of the functional soil with its microbial community did not, over 65 days, revitalize the poorly functioning soil. The findings showed that abiotic limitations to enzymatic activity in barren brownfield soils could be mitigated by establishing primary production but not by the addition of enzymatically active microbial communities alone.

Environmental forensic characterization of former rail yard soils located adjacent to the Statue of Liberty in the New York/New Jersey harbor.

Identifying inorganic and organic soil contaminants in urban brownfields can give insights into the adverse effects of industrial activities on soil function, ecological health, and environmental quality. Liberty State Park in Jersey City (N.J., USA) once supported a major rail yard that had dock facilities for both cargo and passenger service; a portion remains closed to the public, and a forest developed and spread in this area. The objectives of this study were to: 1) characterize the organic and inorganic compounds in Liberty State Park soils and compare the findings to an uncontaminated reference site (Hutcheson Memorial Forest); and 2) identify differences between the barren low-functioning areas and the forested high-functioning areas of the brownfield. Soil samples were solvent-extracted, fractionated, and analyzed by gas chromatography-mass spectrometry and subjected to loss-on-ignition, pyrolysis-gas chromatography-mass spectrometry, inductively-coupled-plasma mass spectrometry, and optical microscopy analyses. Compared to soil from the reference site, the forested soils in Liberty State Park contained elevated percentages of organic matter (30-45%) and more contaminants, such as fossil-fuel-derived hydrocarbons and coal particles. Microscopy revealed bituminous and anthracite coal, coke, tar/pitch, and ash particles. Barren and low-functioning site 25R had a similar organic contaminant profile but contained a higher metal load than other Liberty State Park sites and also lacked higher plant indicators. These can obscure the signatures of contaminants, and data from adjacent barren and vegetated sites are valuable references for soils studies. A deeper understanding of the chemistry, biochemistry, and ecology of barren soils can be leveraged to prevent land degradation and to restore dysfunctional and phytotoxic soils.

Soil microbial response to metal contamination in a vegetated and urban brownfield.

The ubiquity of urban brownfields presents not only a challenge for environmental managers but also an opportunity to study the functional aspects of degraded ecosystems that are in close contact with human habitation. In this study, we investigate the soil microbial community response to heavy metal contamination at Liberty State Park (LSP), an urban brownfield in Jersey City, NJ, USA. Heavy metal contamination of the soils at LSP is heterogeneous, varying widely across site and among metals. We collected soils along a previously mapped gradient of metal contamination at LSP and sampled soil from a local and uncontaminated reference site (Hutcheson Memorial Forest (HMF)) for comparison. For all soils, we measured soil heavy metal concentrations, soil organic carbon content, bacterial density, and extracellular phosphatase activity as a proxy of ecosystem functioning. Additionally, we analyzed the microbial community composition using high-throughput sequencing. Data show that some sites within LSP have significantly higher phosphatase activity compared to HMF, indicating that some heavily contaminated LSP soils are highly functional. We also found that soil organic carbon and bacterial density have a significant and positive relationship with phosphatase activity. The microbial community analyses showed that the bacterial communities were sensitive to heavy metals and that the composition was significantly affected in particular by copper, zinc, and lead. The fungal communities, however, did not vary significantly with heavy metals. Our results shed important light on the composition and functioning of urban brownfield soils. A deeper understanding of these unique ecosystems is required for successful remediation, restoration and urban sustainability.

Abiotic factors determine functional outcomes of microbial inoculation of soils from a metal contaminated brownfield.

Whole community microbial inoculation can improve soil function in contaminated environments. Here we conducted a case study to investigate whether biotic factors (inoculum) or abiotic factors (soil base) have more impact on the extracellular enzymatic activities in a whole community microbial inoculation. To this end, we cross-inoculated microbial communities between two heavy metal-contaminated soils, with high and low extracellular enzyme activities, respectively. We measured extracellular phosphatase activity, a proxy for soil function, after self- and cross-inoculation of microbial communities into sterilized soils, and all activities were normalized to non-inoculated controls. We found that inoculation increased phosphatase activity in the soils. For soils treated with different inocula, we found significant differences in the microbial community compositions but no significant differences in the extracellular phosphatase activities normalized to their respective sterilized, non-inoculated controls (4.7 ±â€¯1.8 and 3.3 ±â€¯0.5 for soils inoculated with microbial communities from 146 to 43, respectively). On the other hand, normalized phosphatase activities between the two soil bases were significantly different (4.1 ±â€¯0.12 and 1.9 ±â€¯0.12 for soil bases 146 and 43, respectively) regardless of the source of the inoculum that did not vary between soil bases. The results indicate that the abiotic properties of the soils were a significant predictor for phosphatase activity but not for the end-point composition of the microbial community. The findings suggest that targeted microbial inocula from metal contaminated soils can increase phosphatase activity, and likely soil functioning in general, but the degree to which this happens depends on the abiotic environment, in this case, metal contamination.

Herbivory and Stoichiometric Feedbacks to Primary Production.

Established theory addresses the idea that herbivory can have positive feedbacks on nutrient flow to plants. Positive feedbacks likely emerge from a greater availability of organic carbon that primes the soil by supporting nutrient turnover through consumer and especially microbially-mediated metabolism in the detrital pool. We developed an entirely novel stoichiometric model that demonstrates the mechanism of a positive feedback. In particular, we show that sloppy or partial feeding by herbivores increases detrital carbon and nitrogen allowing for greater nitrogen mineralization and nutritive feedback to plants. The model consists of differential equations coupling flows among pools of: plants, herbivores, detrital carbon and nitrogen, and inorganic nitrogen. We test the effects of different levels of herbivore grazing completion and of the stoichiometric quality (carbon to nitrogen ratio, C:N) of the host plant. Our model analyses show that partial feeding and plant C:N interact because when herbivores are sloppy and plant biomass is diverted to the detrital pool, more mineral nitrogen is available to plants because of the stoichiometric difference between the organisms in the detrital pool and the herbivore. This model helps to identify how herbivory may feedback positively on primary production, and it mechanistically connects direct and indirect feedbacks from soil to plant production.

Species richness and allometric scaling jointly determine biomass in model aquatic food webs.

1. Allometric theory makes specific predictions about how density, and consequently biomass, scale with organism size within trophic levels, across trophic levels and across food webs. 2. Diversity-yield relationships suggest that more diverse food webs can sometimes support more biomass through mechanisms involving niche complementarity or selection effects that are sometimes attributed to organism size. 3. We combine the above two approaches and show that, generally, density and biomass scale with organism size within and between trophic levels as predicted by allometric theory. Further, food webs converged in total biomass despite persistent differences in the composition and size of the organisms among food webs; species richness explained deviations from the constant yield of biomass expected from size-abundance relationships. 4. Our results suggest that organism size plays only a transient role in controlling community biomass because population increases or decreases lead to rapid convergence in biomass. Species richness affects community biomass independently by effectively increasing the mass of organisms that can be supported in a given productivity regime.

Microbial biogeography: putting microorganisms on the map.

We review the biogeography of microorganisms in light of the biogeography of macroorganisms. A large body of research supports the idea that free-living microbial taxa exhibit biogeographic patterns. Current evidence confirms that, as proposed by the Baas-Becking hypothesis, 'the environment selects' and is, in part, responsible for spatial variation in microbial diversity. However, recent studies also dispute the idea that 'everything is everywhere'. We also consider how the processes that generate and maintain biogeographic patterns in macroorganisms could operate in the microbial world.