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Soil carbon sequestration arises from the interplay of carbon input and stabilization, which vary in space and time. Assessing the resulting microscale carbon distribution in an intact pore space, however, has so far eluded methodological accessibility. Here, we explore the role of soil moisture regimes in shaping microscale carbon gradients by a novel mapping protocol for particulate organic matter and carbon in the soil matrix based on a combination of Osmium staining, X-ray computed tomography, and machine learning. With three different soil types we show that the moisture regime governs C losses from particulate organic matter and the microscale carbon redistribution and stabilization patterns in the soil matrix. Carbon depletion around pores (aperture > 10 µm) occurs in a much larger soil volume (19–74%) than carbon enrichment around particulate organic matter (1%). Thus, interacting microscale processes shaped by the moisture regime are a decisive factor for overall soil carbon persistence. Carbon sequestration in soils has rarely been observed at microscopic scales. Here the authors reveal the impact of soil moisture regimes in shaping carbon stabilization and mineralization patterns tied to the pore network.

Most plants living in tropical acid soils depend on the arbuscular mycorrhizal (AM) symbiosis for mobilizing low-accessible phosphorus (P), due to its strong bonding by iron (Fe) oxides. The roots release low-molecular-weight organic acids (LMWOAs) as a mechanism to increase soil P availability by ligand exchange or dissolution. However, little is known on the LMWOA production by AM fungi (AMF), since most studies conducted on AM plants do not discriminate on the LMWOA origin. This study aimed to determine whether AMF release significant amounts of LMWOAs to liberate P bound to Fe oxides, which is otherwise unavailable for the plant. Solanum lycopersicum L. plants mycorrhized with Rhizophagus irregularis were placed in a bicompartmental mesocosm, with P sources only accessible by AMF. Fingerprinting of LMWOAs in compartments containing free and goethite-bound orthophosphate (OP or GOE-OP) and phytic acid (PA or GOE-PA) was done. To assess P mobilization via AM symbiosis, P content, photosynthesis, and the degree of mycorrhization were determined in the plant; whereas, AM hyphae abundance was determined using lipid biomarkers. The results showing a higher shoot P content, along with a lower N:P ratio and a higher photosynthetic capacity, may be indicative of a higher photosynthetic P-use efficiency, when AM plants mobilized P from less-accessible sources. The presence of mono-, di-, and tricarboxylic LMWOAs in compartments containing OP or GOE-OP and phytic acid (PA or GOE-PA) points toward the occurrence of reductive dissolution and ligand exchange/dissolution reactions. Furthermore, hyphae grown in goethite loaded with OP and PA exhibited an increased content of unsaturated lipids, pointing to an increased membrane fluidity in order to maintain optimal hyphal functionality and facilitate the incorporation of P. Our results underpin the centrality of AM symbiosis in soil biogeochemical processes, by highlighting the ability of the AMF and accompanying microbiota in releasing significant amounts of LMWOAs to mobilize P bound to Fe oxides.

Formation of mineral-organic associations is a key process in the global carbon cycle. Recent concepts propose litter quality-controlled microbial assimilation and direct sorption processes as main factors in transferring carbon from plant litter into mineral-organic associations. We explored the pathways of the formation of mineral-associated organic matter (MOM) in soil profiles along a 120-ky ecosystem gradient that developed under humid climate from the retreating Franz Josef Glacier in New Zealand. We determined the stocks of particulate and mineral-associated carbon, the isotope signature and microbial decomposability of organic matter, and plant and microbial biomarkers (lignin phenols, amino sugars and acids) in MOM. Results revealed that litter quality had little effect on the accumulation of mineral-associated carbon and that plant-derived carbon bypassed microbial assimilation at all soil depths. Seemingly, MOM forms by sorption of microbial as well as plant-derived compounds to minerals. The MOM in carbon-saturated topsoil was characterized by the steady exchange of older for recent carbon, while subsoil MOM arises from retention of organic matter transported with percolating water. Overall, MOM formation is not monocausal but involves various mechanisms and processes, with reactive minerals being effective filters capable of erasing chemical differences in organic matter inputs.

Clay minerals and pedogenic metal (oxyhydr)oxides are the most reactive soil mineral constituents controlling the long-term persistence of organic carbon (OC) in terrestrial ecosystems. However, their co-occurrence in most soils complicates direct assessment of their individual contribution to OC persistence. Making use of unique mineralogical combinations in soils located in the East Usambara Mountains of Tanzania, we disentangled the contribution of clay-sized aluminous minerals (kaolinite, gibbsite) and pedogenic Fe (oxyhydr)oxides (predominant goethite and hematite) on OC storage and stabilization under natural forests and croplands. Topsoil samples, varying in contents but not types of aluminous clays and pedogenic Fe (oxyhydr)oxides, were identified by selective extractions, X-ray diffraction, and Mössbauer spectroscopy. Associated abundance of particulate and mineral-associated organic matter (OM) was quantified by density fractionation and their changes during land-use conversion were determined as a measure of OC persistence. Additionally, we assessed the resistance of OC to chemical oxidation as well as microbial decomposition in a 50-day laboratory incubation. We found that the ratio of pedogenic Fe to aluminous clay is more consequential for OC storage and stabilization than their individual contents, despite the fact that Fe (oxyhydr)oxides generally exert a stronger impact on OC than aluminous clays. Conjunction of large amounts of Fe (oxyhydr)oxides with low aluminous clay contents caused the strongest accumulation of mineral-associated OC, a low soil respiration, high OC stability against chemical oxidation, and high OC persistence during land-use change. Our study suggests that certain mineralogical combinations in the humid tropics alleviate OM losses during land conversion because of the strong and selective mineral control on OC stabilization, particular if the weight ratio of pedogenic Fe to aluminous clay exceeds the threshold range of 0.44‒0.56.

Formation of mineral-associated organic matter (MAOM) is a key process in the global carbon cycle, stabilising organic carbon in soils. The relative importance of mineral composition and land use as potential controls of MAOM stability at regional scales and underlying microbial processes are still unresolved. Here, we assessed the stability of MAOM formed on goethite (iron oxide) and illite (phyllosilicate clay) exposed for five years in topsoils at 68 forest and grassland sites across Germany. We incubated the newly formed MAOM, determined its extractability, and analysed the composition and functioning of associated microbial communities. Decomposition of MAOM was always significantly lower for goethite than illite, highlighting that higher organic carbon accumulation on goethite was not exclusively due to its larger sorption capacity. Instead, reduced organic carbon extractability and higher phosphorus-acquiring enzyme activities indicated stronger substrate limitation of microbial growth on goethite than illite. Across the two minerals, MAOM decomposition was consistently lower for forests than grasslands, relating to greater nutrient constraints and a different microbial community composition in forests. Overall, mineral type and land use explained 34.6 and 23.2% of the variance in MAOM decomposition. The pronounced land use effect on MAOM stability underlines its potential responsiveness to environmental change. Both quantity of organic carbon available to microorganisms and presence of available nutrients control the decomposition of newly formed mineral-associated organic matter, according to a study exposing pristine goethite and illite in topsoils at 68 forest and grassland sites across Germany.

Mangroves play an important role in carbon sequestration, but soil organic carbon (SOC) stocks differ between marine and estuarine mangroves, suggesting differing processes and drivers of SOC accumulation. Here, we compared undegraded and degraded marine and estuarine mangroves in a regional approach across the Indonesian archipelago for their SOC stocks and evaluated possible drivers imposed by nutrient limitations along the land‐to‐sea gradients. SOC stocks in natural marine mangroves (271–572 Mg ha−1 m−1) were much higher than under estuarine mangroves (100–315 Mg ha−1 m−1) with a further decrease caused by degradation to 80–132 Mg ha−1 m−1. Soils differed in C/N ratio (marine: 29–64; estuarine: 9–28), δ15N (marine: −0.6 to 0.7‰; estuarine: 2.5 to 7.2‰), and plant‐available P (marine: 2.3–6.3 mg kg−1; estuarine: 0.16–1.8 mg kg−1). We found N and P supply of sea‐oriented mangroves primarily met by dominating symbiotic N2 fixation from air and P import from sea, while mangroves on the landward gradient increasingly covered their demand in N and P from allochthonous sources and SOM recycling. Pioneer plants favored by degradation further increased nutrient recycling from soil resulting in smaller SOC stocks in the topsoil. These processes explained the differences in SOC stocks along the land‐to‐sea gradient in each mangrove type as well as the SOC stock differences observed between estuarine and marine mangrove ecosystems. This first large‐scale evaluation of drivers of SOC stocks under mangroves thus suggests a continuum in mangrove functioning across scales and ecotypes and additionally provides viable proxies for carbon stock estimations in PES or REDD schemes. In a comparative study of mangrove ecosystem across the Indonesian archipelago, we found SOC stocks in natural marine (271–572 Mg ha−1 m−1) mangroves much higher than under estuarine mangroves (100–315 Mg ha−1 m−1). Soils differed in C/N ratio (marine: 29–64; estuarine: 9–28), δ15N (marine: −0.6 to 0.7‰; estuarine: 2.5 to 7.2‰), and plant‐available P (marine: 2.3–6.3 mg kg−1; estuarine: 0.16–1.8 mg kg−1). N and P supply of sea‐oriented mangroves was primarily met by dominating symbiotic N2 fixation from air and P import from sea, while mangroves on the landward gradient increasingly covered their demand in N and P via SOM recycling. These processes explained the differences in SOC stocks along the land‐to‐sea gradient in each mangrove type as well as the SOC stock differences observed between estuarine and marine mangrove ecosystems.

Adsorption of extracellular enzymes to soil minerals is assumed to protect them against degradation, while modifying their activities at the same time. However, the persistence of the activity of adsorbed enzymes remains poorly understood. Therefore, we studied the persistence of cellulase and α-amylase activities after adsorption to soil amended with various amounts (+1, +5, and +10 wt.%) of three typical soil minerals, montmorillonite, kaolinite, and goethite. Soil without mineral addition (pure soil), pure minerals, and pure dissolved enzymes were used as references. Soil mineral–enzyme complexes were prepared and then incubated for 100 days; temporal changes in enzyme activities were analyzed after 0, 0.1, 1, 10, and 100 days. The specific enzyme activities (activities normalized to protein content) and their persistence (activities relative to activities at day 0) were compared to enzyme activities in solution and after sorption to the control soil. Amylase adsorption to pure minerals increased in the following order: montmorillonite > kaolinite > goethite. That of cellulase increased in the following order: goethite > montmorillonite > kaolinite. Adsorption of enzymes to soils did not increase in the same order of magnitude as the addition of reactive binding sites. Based on inverse relationships between the amount of enzyme adsorbed and the specific enzyme activity and their persistency, we showed that a limited availability of sorption sites is important for high specific activity and persistence of the enzymes. This is probably the consequence of less and weaker bonds, as compared to a high availability of sorption sites, resulting in a smaller impact on the active sites of the enzyme. Hence, we suppose that the soil mineral phase supports microorganisms in less-sorptive environments by saving energy on enzyme production, since small enzyme release could already result in sufficient activities to degrade respective target carbon substrates.

Nutrient supply in phosphorus (P)-limited ecosystems, with most P being associated with secondary minerals, has to rely on efficient nutrient allocation strategies, such as those involving mycorrhizal symbioses. Yet, little is known about the extent of photo-assimilate transfer to the fungal partner, who in turn mobilizes mineral-bound P sources required by the plant. This study aims to explore the carbon (C)–P trade between an arbuscular mycorrhizal (AM) plant and its ability to incorporate P from differently accessible P sources. We compared P uptake rates of AM plants for orthophosphate (OP) and phytic acid (PA), applied to mesocosms in either dissolved form or bound to goethite (α-FeOOH). The design of the mesocosms allowed the plant to only access the P in the fungal compartment via the AM hyphae. We hypothesized the AM plant to invest more C into the symbiosis, if P is present in the less accessible form. To estimate the C budget of the symbiosis, we determined total organic carbon (OC), 16:1ω5c phospholipid fatty acid (PLFA; AM fungi extraradical mycelium), 16:1ω5c neutral lipid fatty acid (NLFA; AM fungi energy storage), and CO2 cumulative respiration in the fungal compartment. A ratio to the total C mobilized into the fungal compartment (OC+CO2 cumulative respiration) and the P incorporated into the AM plant (Total C/P) was calculated to estimate the C investment made by the AM plant into its symbiotic partner. AM plants incorporated P derived from all four P sources exclusively via the mycorrhizal pathway in different amounts and kinetics. The Total C/P ratio was significantly larger for those AM plants accessing the goethite-bound P compounds. They also transferred significantly larger amounts of PLFA and NLFA to their fungal partner, both indicating a larger plant C investment per P taken up. Our data provide first evidence about the ability of an AM plant to incorporate P from an organic source bound to a secondary mineral. The different C investments of AM plants into P allocation from variably available sources suggests a broad nexus between P mining strategies, resource partitioning in soil, and the amounts of C accumulated in terrestrial soils.

Enzyme-mediated decomposition of soil organic matter (SOM) is controlled, amongst other factors, by organic matter properties and by the microbial decomposer community present. Since microbial community composition and SOM properties are often interrelated and both change with soil depth, the drivers of enzymatic decomposition are hard to dissect. We investigated soils from three regions in the Siberian Arctic, where carbon rich topsoil material has been incorporated into the subsoil (cryoturbation). We took advantage of this subduction to test if SOM properties shape microbial community composition, and to identify controls of both on enzyme activities. We found that microbial community composition (estimated by phospholipid fatty acid analysis), was similar in cryoturbated material and in surrounding subsoil, although carbon and nitrogen contents were similar in cryoturbated material and topsoils. This suggests that the microbial community in cryoturbated material was not well adapted to SOM properties. We also measured three potential enzyme activities (cellobiohydrolase, leucine-amino-peptidase and phenoloxidase) and used structural equation models (SEMs) to identify direct and indirect drivers of the three enzyme activities. The models included microbial community composition, carbon and nitrogen contents, clay content, water content, and pH. Models for regular horizons, excluding cryoturbated material, showed that all enzyme activities were mainly controlled by carbon or nitrogen. Microbial community composition had no effect. In contrast, models for cryoturbated material showed that enzyme activities were also related to microbial community composition. The additional control of microbial community composition could have restrained enzyme activities and furthermore decomposition in general. The functional decoupling of SOM properties and microbial community composition might thus be one of the reasons for low decomposition rates and the persistence of 400 Gt carbon stored in cryoturbated material.

Arctic plant productivity is often limited by low soil N availability. This has been attributed to slow breakdown of N-containing polymers in litter and soil organic matter (SOM) into smaller, available units, and to shallow plant rooting constrained by permafrost and high soil moisture. Using 15N pool dilution assays, we here quantified gross amino acid and ammonium production rates in 97 active layer samples from four sites across the Siberian Arctic. We found that amino acid production in organic layers alone exceeded literature-based estimates of maximum plant N uptake 17-fold and therefore reject the hypothesis that arctic plant N limitation results from slow SOM breakdown. High microbial N use efficiency in organic layers rather suggests strong competition of microorganisms and plants in the dominant rooting zone. Deeper horizons showed lower amino acid production rates per volume, but also lower microbial N use efficiency. Permafrost thaw together with soil drainage might facilitate deeper plant rooting and uptake of previously inaccessible subsoil N, and thereby promote plant productivity in arctic ecosystems. We conclude that changes in microbial decomposer activity, microbial N utilization and plant root density with soil depth interactively control N availability for plants in the Arctic.