Nitrification Control by Plants and Preference for Ammonium versus Nitrate: Positive Feedbacks Increase Productivity but Undermine Resilience.

AbstractSome plants, via their action on microorganisms, control soil nitrification (i.e., the transformation of ammonium into nitrate). We model how the covariation between plant control of nitrification and preference for ammonium versus nitrate impacts ecosystem properties such as productivity, nitrogen (N) losses, and overall resilience. We show that the control of nitrification can maximize productivity by minimizing total inorganic N losses. We initially predicted that plants with an ammonium preference should achieve the highest biomass when inhibiting nitrification, and conversely that plants preferring nitrate should achieve the highest biomass by stimulating nitrification. With a parametrization derived from the Lamto savanna (Ivory Coast), we find that productivity is maximal for plants that slightly prefer ammonium and inhibit nitrification. Such situations, however, lead to strong positive feedbacks that can cause abrupt shifts from a highly to a lowly productive ecosystem. The comparison with other parameter sets (Pawnee short-grass prairie [United States], intensively cultivated field, and a hypothetical parameter set in which ammonium is highly volatilized and nitrate inputs are high) shows that strategies yielding the highest biomass may be counterintuitive (i.e., preferring nitrate but inhibiting nitrification). We argue that the level of control yielding the highest productivity depends on ecosystem properties (quantity of N deposition, leaching rates, and baseline nitrification rates), not only preference. Finally, while contrasting N preferences offer, as expected, the possibility of coexistence through niche partitioning, we stress how control of nitrification can be framed as a niche construction process that adds an additional dimension to coexistence conditions.

Short-term impact of fire on the total soil microbial and nitrifier communities in a wet savanna.

Savannas are characterized by the coexistence of grasses and trees. Fires are critical for their coexistence, because they decrease the survival of tree seedlings and saplings and their recruitment to the adult stage. In some humid savannas, perennial grasses inhibit nitrification and trees stimulate nitrification, which likely favors coexistence between trees and grasses. However, fires may influence plant capacity to control nitrogen cycling, which could subsequently influence tree-grass coexistence and savanna nitrogen budget. Therefore, we sampled soil in a humid savanna of Ivory Coast under the dominant nitrification-inhibiting grass species and the dominant nitrification-stimulating tree species and under bare soil before and after (i.e., 5 days) fire during the long dry season. We quantified the total microbial and nitrifier abundances and transcriptional activities and the nitrification enzyme activity. Fire decreased soil water content, probably by increasing evaporation and, maybe, by triggering the growth of grasses, and increased soil ammonium availability likely due to ash deposition and increased mineralization. Fire did not impact the total archaeal, bacterial, or fungal abundances, or that of the nitrifiers. Fire did not impact archaeal transcriptional activity and increased bacterial and fungal total transcriptional activities. In contrast, fire decreased the archaeal nitrifier transcriptional activities and the nitrification enzymatic activity, likely due to the often reported resumption of the growth of nitrification-inhibiting grasses quickly after the fire (and the subsequent increase in root exudation). These results pave the way for a better understanding of the short-term effects of fire on nitrogen cycling and tree-grass competition for nitrogen.

Influence of the residence time of street trees and their soils on trace element contamination in Paris (France).

With the actual increasing interest for urban soils, the evaluation of soil contamination by trace elements and the dynamics of this contamination appear mandatory to preserve plant and thereby human health. Street trees and the associated soil placed in pits located nearby roads could represent convenient indicators of urban and vehicle traffic influences on soils and plants. However, data on these soils remain scarce, many studies investigating park soils rather than street tree soils. Furthermore, trace elements could be one of the main factors causing the observed urban tree decline, while practitioners more and more question the possible reuse of these soils after the death of trees as well as tree litter collected in the streets. We evaluated the contamination in anthropogenic trace elements (TE), namely Zn, Pb, and Cd, of street trees (Tilia tomentosa) and their soils distributed all over Paris (France). Street tree soils are imported from rural areas at the plantation of each new tree so that tree age corresponds to the time of residence of the soil within an urban environment allowing the evaluation of temporal trends on TE concentration in soils and trees. The TE concentration revealed an important soil pollution, especially for the older soils (mean age of 80 years old). The consideration of the residence time of trees and soils in an urban environment evidenced an accumulation of Zn and Pb (ca. 4.5 mg kg-1 year-1 and 4 mg kg-1 year-1 for Zn and Pb, respectively). However, leaf concentrations in TE were low and indicate that soil-root transfer was not significant compared to the contamination by atmospheric deposition. These results underlined the necessity to deepen the evaluation of the recycling of urban soils or plants submitted to urban contamination.

Genotypic variability enhances the reproducibility of an ecological study.

Many scientific disciplines are currently experiencing a 'reproducibility crisis' because numerous scientific findings cannot be repeated consistently. A novel but controversial hypothesis postulates that stringent levels of environmental and biotic standardization in experimental studies reduce reproducibility by amplifying the impacts of laboratory-specific environmental factors not accounted for in study designs. A corollary to this hypothesis is that a deliberate introduction of controlled systematic variability (CSV) in experimental designs may lead to increased reproducibility. To test this hypothesis, we had 14 European laboratories run a simple microcosm experiment using grass (Brachypodium distachyon L.) monocultures and grass and legume (Medicago truncatula Gaertn.) mixtures. Each laboratory introduced environmental and genotypic CSV within and among replicated microcosms established in either growth chambers (with stringent control of environmental conditions) or glasshouses (with more variable environmental conditions). The introduction of genotypic CSV led to 18% lower among-laboratory variability in growth chambers, indicating increased reproducibility, but had no significant effect in glasshouses where reproducibility was generally lower. Environmental CSV had little effect on reproducibility. Although there are multiple causes for the 'reproducibility crisis', deliberately including genetic variability may be a simple solution for increasing the reproducibility of ecological studies performed under stringently controlled environmental conditions.

Trace element concentrations along a gradient of urban pressure in forest and lawn soils of the Paris region (France).

The concentration, degree of contamination and pollution of 7 trace elements (TEs) along an urban pressure gradient were measured in 180 lawn and wood soils of the Paris region (France). Iron (Fe), a major element, was used as reference element. Copper (Cu), cadmium (Cd), lead (Pb) and zinc (Zn) were of anthropogenic origin, while arsenic (As), chromium (Cr) and nickel (Ni) were of natural origin. Road traffic was identified as the main source of anthropogenic TEs. In addition, the industrial activity of the Paris region, especially cement plants, was identified as secondary source of Cd. Soil characteristics (such as texture, organic carbon (OC) and total nitrogen (tot N) contents) tell the story of the soil origins and legacies along the urban pressure gradient and often can explain TE concentrations. The history of the land-use types was identified as a factor that allowed understanding the contamination and pollution by TEs. Urban wood soils were found to be more contaminated and polluted than urban lawns, probably because woods are much older than lawns and because of the legacy of the historical management of soils in the Paris region (Haussmann period). Lawn soils are similar to the fertile agricultural soils and relatively recently (mostly from the 1950s onwards) imported from the surrounding of Paris, so that they may be less influenced by urban conditions in terms of TE concentrations. Urban wood soils are heavily polluted by Cd, posing a high risk to the biological communities. The concentration of anthropogenic TEs increased from the rural to the urban areas, and the concentrations of most anthropogenic TEs in urban areas were equivalent to or above the regulatory reference values, raising the question of longer-term monitoring.

Multifunctionality is affected by interactions between green roof plant species, substrate depth, and substrate type.

Green roofs provide ecosystem services through evapotranspiration and nutrient cycling that depend, among others, on plant species, substrate type, and substrate depth. However, no study has assessed thoroughly how interactions between these factors alter ecosystem functions and multifunctionality of green roofs. We simulated some green roof conditions in a pot experiment. We planted 20 plant species from 10 genera and five families (Asteraceae, Caryophyllaceae, Crassulaceae, Fabaceae, and Poaceae) on two substrate types (natural vs. artificial) and two substrate depths (10 cm vs. 30 cm). As indicators of major ecosystem functions, we measured aboveground and belowground biomasses, foliar nitrogen and carbon content, foliar transpiration, substrate water retention, and dissolved organic carbon and nitrates in leachates. Interactions between substrate type and depth strongly affected ecosystem functions. Biomass production was increased in the artificial substrate and deeper substrates, as was water retention in most cases. In contrast, dissolved organic carbon leaching was higher in the artificial substrates. Except for the Fabaceae species, nitrate leaching was reduced in deep, natural soils. The highest transpiration rates were associated with natural soils. All functions were modulated by plant families or species. Plant effects differed according to the observed function and the type and depth of the substrate. Fabaceae species grown on natural soils had the most noticeable patterns, allowing high biomass production and high water retention but also high nitrate leaching from deep pots. No single combination of factors enhanced simultaneously all studied ecosystem functions, highlighting that soil-plant interactions induce trade-offs between ecosystem functions. Substrate type and depth interactions are major drivers for green roof multifunctionality.

Assessment of trace metal air pollution in Paris using slurry-TXRF analysis on cemetery mosses.

Mosses are useful, ubiquitous accumulation biomonitors and as such can be used for biomonitoring surveys. However, the biomonitoring of atmospheric pollution can be compromised in urban contexts if the targeted biomonitors are regularly disturbed, irregularly distributed, or are difficult to access. Here, we test the hypothesis that cemeteries are appropriate moss sampling sites for the evaluation of air pollution in urban areas. We sampled mosses growing on gravestones in 21 urban and peri-urban cemeteries in the Paris metropolitan area. We focused on Grimmia pulvinata (Hedwig) Smith, a species abundantly found in all studied cemeteries and very common in Europe. The concentration of Al, As, Br, Ca, Ce, Cl, Cr, Cu, Fe, K, Mn, Ni, V, P, Pb, Rb, S, Sr, Ti, and Zn was determined by a total reflection X-ray fluorescence technique coupled with a slurry sampling method (slurry-TXRF). This method avoids a digestion step, reduces the risk of sample contamination, and works even at low sample quantities. Elemental markers of road traffic indicated that the highest polluted cemeteries were located near the highly frequented Parisian ring road and under the influence of prevailing winds. The sites with the lowest pollution were found not only in the peri-urban cemeteries, adjoining forest or farming landscapes, but also in the large and relatively wooded cemeteries located in the center of Paris. Our results suggest that (1) slurry-TXRF might be successfully used with moss material, (2) G. pulvinata might be a good biomonitor of trace metals air pollution in urban context, and (3) cemetery moss sampling could be a useful complement for monitoring urban areas. Graphical abstract We tested the hypothesis that cemeteries are appropriate moss sampling sites for the evaluation of air pollution in urban areas. We sampled 110 moss cushions (Grimmia pulvinata) growing on gravestones in 21 urban and peri-urban cemeteries in the Paris metropolitan area. The concentration of 20 elements in mosses was determined by a total reflection X-ray fluorescence technique coupled with a slurry sampling method. Statistical analysis revealed that: - Urbanized Parisian areas crossed by traffic roads have the highest polluted cemeteries with a strong influence of main wind direction on the distribution of air pollutants - As expected, small cemeteries with low tree density were heavily polluted - Less obvious, large green spaces such as large cemeteries (Père Lachaise, Montmartre, Montparnasse) in the center of a dense metropolis like Paris present the same level of atmospheric trace metal pollution as cemeteries in less urbanized areas or nearing a very large forest. This suggests that even in densely urbanized areas, there is more spatial variability in pollution distribution than usually assumed and that large urban areas with low traffic and green filters such as trees are likely to intercept air pollutants.

Suppression of soil nitrification by plants.

Nitrification, the biological oxidation of ammonium to nitrate, weakens the soil's ability to retain N and facilitates N-losses from production agriculture through nitrate-leaching and denitrification. This process has a profound influence on what form of mineral-N is absorbed, used by plants, and retained in the soil, or lost to the environment, which in turn affects N-cycling, N-use efficiency (NUE) and ecosystem health and services. As reactive-N is often the most limiting in natural ecosystems, plants have acquired a range of mechanisms that suppress soil-nitrifier activity to limit N-losses via N-leaching and denitrification. Plants' ability to produce and release nitrification inhibitors from roots and suppress soil-nitrifier activity is termed 'biological nitrification inhibition' (BNI). With recent developments in methodology for in-situ measurement of nitrification inhibition, it is now possible to characterize BNI function in plants. This review assesses the current status of our understanding of the production and release of biological nitrification inhibitors (BNIs) and their potential in improving NUE in agriculture. A suite of genetic, soil and environmental factors regulate BNI activity in plants. BNI-function can be genetically exploited to improve the BNI-capacity of major food- and feed-crops to develop next-generation production systems with reduced nitrification and N2O emission rates to benefit both agriculture and the environment. The feasibility of such an approach is discussed based on the progresses made.

A tale of four stories: soil ecology, theory, evolution and the publication system.

BACKGROUND: Soil ecology has produced a huge corpus of results on relations between soil organisms, ecosystem processes controlled by these organisms and links between belowground and aboveground processes. However, some soil scientists think that soil ecology is short of modelling and evolutionary approaches and has developed too independently from general ecology. We have tested quantitatively these hypotheses through a bibliographic study (about 23000 articles) comparing soil ecology journals, generalist ecology journals, evolutionary ecology journals and theoretical ecology journals. FINDINGS: We have shown that soil ecology is not well represented in generalist ecology journals and that soil ecologists poorly use modelling and evolutionary approaches. Moreover, the articles published by a typical soil ecology journal (Soil Biology and Biochemistry) are cited by and cite low percentages of articles published in generalist ecology journals, evolutionary ecology journals and theoretical ecology journals. CONCLUSION: This confirms our hypotheses and suggests that soil ecology would benefit from an effort towards modelling and evolutionary approaches. This effort should promote the building of a general conceptual framework for soil ecology and bridges between soil ecology and general ecology. We give some historical reasons for the parsimonious use of modelling and evolutionary approaches by soil ecologists. We finally suggest that a publication system that classifies journals according to their Impact Factors and their level of generality is probably inadequate to integrate "particularity" (empirical observations) and "generality" (general theories), which is the goal of all natural sciences. Such a system might also be particularly detrimental to the development of a science such as ecology that is intrinsically multidisciplinary.