Demand of all living organisms on the same nutrients forms the basis for interspecific competition between plants and microorganisms in soils. This competition is especially strong in the rhizosphere. To evaluate competitive and mutualistic interactions between plants and microorganisms and to analyse ecological consequences of these interactions, we analysed 424 data pairs from 41 (15)N-labelling studies that investigated (15)N redistribution between roots and microorganisms. Calculated Michaelis-Menten kinetics based on K(m) (Michaelis constant) and V(max) (maximum uptake capacity) values from 77 studies on the uptake of nitrate, ammonia, and amino acids by roots and microorganisms clearly showed that, shortly after nitrogen (N) mobilization from soil organic matter and litter, microorganisms take up most N. Lower K(m) values of microorganisms suggest that they are especially efficient at low N concentrations, but can also acquire more N at higher N concentrations (V(max)) compared with roots. Because of the unidirectional flow of nutrients from soil to roots, plants are the winners for N acquisition in the long run. Therefore, despite strong competition between roots and microorganisms for N, a temporal niche differentiation reflecting their generation times leads to mutualistic relationships in the rhizosphere. This temporal niche differentiation is highly relevant ecologically because it: protects ecosystems from N losses by leaching during periods of slow or no root uptake; continuously provides roots with available N according to plant demand; and contributes to the evolutionary development of mutualistic interactions between roots and microorganisms.

Competition between roots and microorganisms for nitrogen: mechanisms and ecological relevance

Predicting and mitigating changes in soil carbon (C) stocks under global change requires a coherent understanding of the factors regulating soil organic matter (SOM) formation and persistence, including knowledge of the direct sources of SOM (plants vs. microbes). In recent years, conceptual models of SOM formation have emphasized the primacy of microbial‐derived organic matter inputs, proposing that microbial physiological traits (e.g., growth efficiency) are dominant controls on SOM quantity. However, recent quantitative studies have challenged this view, suggesting that plants make larger direct contributions to SOM than is currently recognized by this paradigm. In this review, we attempt to reconcile these perspectives by highlighting that variation across estimates of plant‐ versus microbial‐derived SOM may arise in part from methodological limitations. We show that all major methods used to estimate plant versus microbial contributions to SOM have substantial shortcomings, highlighting the uncertainty in our current quantitative estimates. We demonstrate that there is significant overlap in the chemical signatures of compounds produced by microbes, plant roots, and through the extracellular decomposition of plant litter, which introduces uncertainty into the use of common biomarkers for parsing plant‐ and microbial‐derived SOM, especially in the mineral‐associated organic matter (MAOM) fraction. Although the studies that we review have contributed to a deeper understanding of microbial contributions to SOM, limitations with current methods constrain quantitative estimates. In light of recent advances, we suggest that now is a critical time to re‐evaluate long‐standing methods, clearly define their limitations, and develop a strategic plan for improving the quantification of plant‐ and microbial‐derived SOM. From our synthesis, we outline key questions and challenges for future research on the mechanisms of SOM formation and stabilization from plant and microbial pathways.

Clarifying the evidence for microbial‐ and plant‐derived soil organic matter, and the path toward a more quantitative understanding

Elucidating the processes underlying the persistence of soil organic matter (SOM) is a prerequisite for projecting soil carbon feedback to climate change. However, the potential role of plant carbon input in regulating the multi-layer SOM preservation over broad geographic scales remains unclear. Based on large-scale soil radiocarbon (∆14 C) measurements on the Tibetan Plateau, we found that plant carbon input was the major contributor to topsoil carbon destabilisation despite the significant associations of topsoil ∆14 C with climatic and mineral variables as well as SOM chemical composition. By contrast, mineral protection by iron-aluminium oxides and cations became more important in preserving SOM in deep soils. These regional observations were confirmed by a global synthesis derived from the International Soil Radiocarbon Database (ISRaD). Our findings illustrate different effects of plant carbon input on SOM persistence across soil layers, providing new insights for models to better predict multi-layer soil carbon dynamics under changing environments.

Soil carbon persistence governed by plant input and mineral protection at regional and global scales

Despite the fact that heterotrophic nitrification was identified more than 100 years ago, the biochemistry of heterotrophic nitrifiers is poorly known and their contribution to nitrification in soil is still speculative. Heterotrophic nitrifiers need organic compounds as their energy source in contrast to the chemolithotrophic nitrifiers. Most of the potential pathways for nitrite/nitrate production by heterotrophs can be considered as secondary metabolism. Only nitrification and simultaneous denitrification by some heterotrophic bacteria is known to have connection to the energy metabolism. Evidently, the nitrification pathways of bacteria and fungi differ. Some heterotrophic bacteria oxidizing ammonia have ammonia monooxygenase (AMO), but there are also heterotrophic bacteria oxidizing ammonia without AMO. The structure of AMO of chemolithotrophic ammonia oxidizers and heterotrophs differs. AMO has not been found in nitrifying fungi. The conditions for heterotrophic nitrification in soil highly differ from those in waste waters where heterotrophic nitrification activity can be high. Heterotrophic nitrification in soil is limited by the low availability of easily decomposable organic substrates. Possible nitrification by heterotrophs in the rhizosphere and endophytic root microbes gaining a good supply of substrates from plants is not known. Fungi are of special interest in heterotrophic nitrification in soil because fungi can evidently nitrify not only easily decomposable substrates but also when decomposing recalcitrant organic compounds (like lignin) which are abundant in soil. Nitrite/nitrate production from easily decomposable nitrogenous organic compounds, such as amino acids, is common among heterotrophic bacteria and fungi. The general conclusion that heterotrophs use solely an “organic pathway” in their nitrification is, however, not valid because also ammonia can be oxidized. Owing to the diverse, poorly known biochemistry of heterotrophic nitrification and methodological difficulties to differentiate heterotrophic and chemolithotrophic nitrification in soil, the role of heterotrophic nitrification in soil nitrogen cycle remains uncertain. • Heterotrophic nitrification involves several poorly known mechanisms. • Heterotrophic bacteria, unlike fungi can possess ammonia monooxygenase. • Fungi are able to nitrify easily decomposable and recalcitrant organic compounds. • The methods available do not allow accurate determination of nitrification in soil. 

Heterotrophic nitrification – An eternal mystery in the nitrogen cycle

Global atmospheric nitrogen deposition significantly affects the nutrient cycling and C–N–P stoichiometry in ecosystems. Herein, a global meta-analysis was conducted based on 898 pairwise observations to analyze the impact of nitrogen addition on plant-soil-enzyme C–N–P stoichiometry and microbial nutrient limitation in different ecosystem types (cropland, grassland, and forest), nitrogen addition intensity (0–5, 5–10, and >10 g N m−2 yr−1) and duration (0–5, 5–10, and >10yr). Results showed that nitrogen addition significantly decreased plant C:N (shoot: 16.5%, root: 27.1%, litter: 16.5%), soil C:N (5.9%), enzyme C:P (1.2%), and enzyme N:P (5.1%), whereas significantly increased soil C:P (4.9%), enzyme C:N (7.1%), vector angle (4.4%), vector length (3.9%), and plant N:P (shoot: 24.1%, root: 23.8%, and litter: 13.5%). Furthermore, nitrogen addition mainly affected the enzyme C:N and vector length in grasslands. Additionally, the changes in C:N in plants, soil, and enzymes, and vector angle and length were higher at nitrogen addition intensity of >10 g N m−2 yr−1. The changes in C:N and C:P in plant and soil were higher at nitrogen addition duration of >10 yr. Finally, the N:P in shoot, soil and enzyme, and vector angle were strongly correlated with mean annual precipitation (MAP). In conclusion, nitrogen addition significantly reduced the C:N ratio in plants and soil and increased plant N:P, and microbial C and P limitation. These effects vary with the ecosystem type, MAP, and nitrogen addition intensity and duration. The results improve our understanding of the plant-soil-microbial nutrient cycling processes in terrestrial ecosystems under global nitrogen deposition. 

Impact of nitrogen addition on plant-soil-enzyme C–N–P stoichiometry and microbial nutrient limitation

Key Findings: Combining physical fractionation and pyrolysis–gas chromatography/mass spectrometry (py-GC/MS) technique can help better understand the dynamics of soil organic matter (SOM). Background and Objectives: SOM plays a critical role in the global carbon (C) cycle. However, its complexity remains a challenge in characterizing chemical molecular composition within SOM and under nitrogen (N) deposition. Materials and Methods: Three particulate organic matter (POM) fractions within SOM and under N treatments were studied from perspectives of distributions, C contents and chemical signatures in a subtropical forest. N addition experiment was conducted with two inorganic N forms (NH4Cl and NaNO3) applied at three rates of 0, 40, 120 kg N ha−1 yr−1. Three particle-size fractions (>250 μm, 53–250 μm and <53 μm) were separated by a wet-sieving method. Py-GC/MS technique was used to differentiate between chemical composition. Results: A progressive proportion transfer of mineral-associated organic matter (MAOM) to fine POM under N treatment was found. Only C content in fine POM was sensitive to N addition. Principal component analyses (PCA) showed that the coarse POM had the largest plant-derived markers (lignins, phenols, long-chain n-alkanes, and n-alkenes). Short-chain n-alkanes and n-alkenes, benzofurans, aromatics and polycyclic aromatic hydrocarbons mainly from black carbon prevailed in the fine POM. N compounds and polysaccharides from microbial products dominated in the MAOM. Factor analysis revealed that the degradation extent of three fractions was largely distinct. The difference in chemical structure among three particulate fractions within SOM was larger than treatments between control and N addition. In terms of N treatment impact, the MAOM fraction had fewer benzofurans compounds and was enriched in polysaccharides, indicating comparatively weaker mineralization and stronger stabilization of these substances. Conclusions: Our findings highlight the importance of chemical structure in SOM pools and help to understand the influence of N deposition on SOM transformation.

/pdf/different-molecular-characterization-of-soil-particulate-4h697isevm.pdf

Different Molecular Characterization of Soil Particulate Fractions under N Deposition in a Subtropical Forest

Organic nitrogen addition causes decoupling of microbial nitrogen cycles by stimulating gross nitrogen transformation in a temperate forest soil

Feammox is more important than anammox in anaerobic ammonium loss in farmland soils around Lake Taihu, China.

Alteration in enzymatic stoichiometry controls the response of soil organic carbon dynamic to nitrogen and water addition in temperate cultivated grassland

Exogenous nitrogen (N) inputs greatly change the emission and uptake of carbon dioxide (CO2) and methane (CH4) from temperate grassland soils, thereby affecting the carbon (C) budget of regional terrestrial ecosystems. Relevant research focused on natural grassland, but the effects of N fertilization on C exchange fluxes from different forage soils and the driving mechanisms were poorly understood. Here, a three-year N addition experiment was conducted on cultivated grassland planted with alfalfa (Medicago sativa) and bromegrass (Bromus inermis) in Inner Mongolia. The fluxes of soil-atmospheric CO2 and CH4; the content of the total dissolved N (TDN); the dissolved organic N (DON); the dissolved organic C (DOC); NH4+–N and NO3−–N in soil; enzyme activity; and auxiliary variables (soil temperature and moisture) were simultaneously measured. The results showed that N fertilization (>75 kg N ha−1 year−1) caused more serious soil acidification for alfalfa planting than for brome planting. N fertilization stimulated P-acquiring hydrolase (AP) in soil for growing Bromus inermis but did not affect C- and N-acquiring hydrolases (AG, BG, CBH, BX, LAP, and NAG). The oxidase activities (PHO and PER) of soil for planting Bromus inermis were higher than soil for planting Medicago sativa, regardless of N, whether fertilization was applied or not. Forage species and N fertilization did not affect soil CO2 flux, whereas a high rate of N fertilization (150 kg N ha−1 year−1) significantly inhibited CH4 uptake in soil for planting Medicago sativa. A synergistic effect between CO2 emission and CH4 uptake in soil was found over the short term. Our findings highlight that forage species affect soil enzyme activity in response to N fertilization. Soil enzyme activity may be an important regulatory factor for C exchange from temperate artificial grassland soil in response to N fertilization.

Soil Enzyme Activity Regulates the Response of Soil C Fluxes to N Fertilization in a Temperate Cultivated Grassland

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