Fungal Necromass Research Articles

Arbuscular mycorrhizal symbiosis enhances microbial contribution to mineral-associated organic carbon persistence in the soil: Insights from soil microbial community and microbial necromass carbon

The formation of mineral-associated organic carbon (MAOC), a critical soil fraction related to soil organic carbon (SOC) sequestration, is significantly influenced by microbial processes. Yet, how arbuscular mycorrhizal (AM) symbiosis impacts MAOC accumulation in rhizosphere soils through its effects on microbial contribution remains poorly understood. Here, we assessed the microbial community of different soil samples using 16S rRNA sequencing and found that AM symbiosis influenced the soil microbial community composition and enhanced soil microbial community functions related to carbon (C) degradation. The numbers of live and dead cells (all cells in the soil) were highest in AM-RS (rhizosphere soils in the AMF treatment) based on confocal laser scanning microscopy (CLSM) images. Meanwhile, AM symbiosis increased the proportions of apoptotic cells (19.02%) and necrotic cells (12.12%) in rhizosphere soils. Using soil amino sugars as microbial biomarkers, the concentrations of bacterial necromass C (188.01 g kg-1 OC) and fungal necromass C (392.19 g kg-1 OC) were highest in the MAOC fraction in AM-RS. Additionally, the MAOC content (17.20 g kg-1 soil) and proportion (42.06%) were highest in AM-RS. This research illustrates two primary mechanisms by which AM symbiosis enhances MAOC accumulation: (1) by altering the microbial community composition and function related to C degradation; and (2) by promoting the input of microbial necromass C, especially fungal necromass C. This research broadens our horizons to understand the mechanism of microbial contribution to MAOC accumulation stimulated by AM symbiosis in rhizosphere soils and provides management practices for AMF application to SOC sequestration.

Characterization of organic materials in regulating soil carbon sequestration of urban greenspaces: Links to microbial necromass and community composition

Application of organic material is a win-win strategy that effectively boosts soil carbon (C) storage and promotes organic waste recycling in urban ecosystems. However, the divergent responses of C sequestration to organic materials and the underlying mechanisms remain unclear. Using molecular and litterbag methods, this study examined soil organic C (SOC) content, C sequestration efficiency of organic material (CSE-organic material), microbial necromass and community composition in urban greenspaces amended with different types of organic materials. The field experiment had seven treatments: addition of green waste (GreenWaste), green waste compost (GreenWasteCompost), biogas residue (BiogasResidue), biogas residue compost (BiogasResidueCompost), peat (PEAT), biochar (BioChar), and no organic material (Control). Organic materials after 16 months application increased SOC content by 34.1–87.0% compared with the Control and presented CSE-organic material in the order: BioChar > GreenWasteCompost > PEAT > BiogasResidueCompost > GreenWaste > BiogasResidue. Aromaticity index was positively correlated with CSE-organic material, indicating that aromatic C addition had a strong capacity to enhance C sequestration. Microbial necromass increased from 2.7 g C kg−1 in the Control to 3.9–5.0 g C kg−1 under organic materials addition. Bacterial necromass was enriched in the BiogasResidue and BiogasResidueCompost treatments, primarily because of sufficient substrate that facilitated the proliferation and activity of copiotrophic Firmicutes, whereas fungal necromass was 8.8–19.1% higher in the GreenWaste, GreenWasteCompost, and PEAT treatments than that in the BiogasResidue and BiogasResidueCompost treatments, accompanying by an increasing abundance of Ascomycota. However, the contribution of microbial necromass to the increased SOC was negatively correlated with CSE-organic material, suggesting that the input of recalcitrant C weakened the role of microbial necromass in C retention. Overall, this study reveals that the efficiency of organic materials application in promoting soil C accumulation depends largely on recalcitrant C rather than microbial necromass, which highlights the important role of organic waste acting as a C source in achieving urban sustainable development.

Microbial necromass and glycoproteins for determining soil carbon formation under arbuscular mycorrhiza symbiosis

Arbuscular mycorrhizal fungi (AMF) form symbioses with most terrestrial plants and critically modulate soil organic carbon (C) dynamics. Whether AMF promote soil C storage and stability is, however, largely unknown. Since microbial necromass C (MNC) and glomalin-related soil protein (GRSP) are stable microbial-derived C in soils, we therefore evaluated how AMF symbiosis alters both soil C pools and their contributions to soil organic C (SOC) under nitrogen fertilization, based on a 16-weeks mesocosm experiment using a mutant tomato with highly reduced AMF symbiosis. Results showed that SOC content is 4.5 % higher following AMF symbiosis. Additionally, the content of MNC and total GRSP were 47.5 % and 22.3 % higher under AMF symbiosis than at AMF absence, respectively. The accumulations of GRSP and microbial necromass in soil were closely associated with mineral-associated organic C and the abundance of AMF. The increased soil living microbial biomass under AMF symbiosis was mainly derived from AMF biomass, and fungal necromass C significantly contributed to SOC accumulation, as evidenced by the higher fungal:bacterial necromass C ratio under AMF symbiosis. On the contrary, bacterial necromass was degraded to compensate for the increased microbial nutrient demand because of the aggravated nutrient limitation under AMF symbiosis, leading to a decrease in bacterial necromass. Redundancy analysis showing that bacterial necromass was negatively correlated with soil C:N ratio supported this argument. Moreover, the relative change rate of total GRSP was consistently greater in nitrogen-limited soil than that of microbial necromass. Our findings suggested GRSP accumulates faster and contributes more to SOC pools under AMF symbiosis than microbial necromass. The positive correlation between the contributions of GRSP and MNC to SOC further provided valuable information in terms of enhancing our understanding of mechanisms underlying the maintenance of SOC stocks through microbial-derived C.

Necromass responses to warming: A faster microbial turnover in favor of soil carbon stabilisation

Microbial byproducts and residues (hereafter ‘necromass’) potentially play the most critical role in soil organic carbon (SOC) sequestration. However, little is known about the influence of climate warming on necromass accumulation in the agroecosystem and the underlying mechanisms associated with microbial life strategies. In order to address these knowledge gaps, we used amino sugars as biomarkers of microbial necromass, and investigated their variation through an 8-year trial in an agroecosystem with two warming levels (+1.6 and + 3.2 °C) compared to ambient temperature. The results showed that the lower warming level had no impact on total microbial necromass carbon. Conversely, warming the soil 3.2 °C above ambient increased total microbial necromass by 17 % and its contribution to SOC by 21.3 %, mainly by increasing fungal necromass (+19.8 %), whereas +3.2 °C warming had no impact on bacterial necromass. At the phylum level, compared with the ambient control, +3.2 °C warming induced an increase in the abundance of Proteobacteria and a decrease in both Acidobacteria and Actinobacteria, whereas in the fungal community, Ascomycota increased and Mortierellomycota decreased. This indicates that r-strategists outcompete K-strategists in warmer climates, which led to increased microbial necromass production and accumulation, as supported by the positive correlation between r-strategists and microbial necromass. Stronger microbial competition for resources also resulted in a higher biomass turnover rate, greater cell death, and greater production of microbial necromass. This was supported by the lower bacterial and fungal network complexity and trophic links under warming conditions. In addition, the necromass generated from accelerated microbial turnover further offsets warming-induced deceases in microbial biomass. Consequently, bulk SOC did not change, despite microbial necromass having a much greater response to warming than the soil C pool. Therefore, future climate warming may influence the composition and persistence of SOC during microbial degradation.

Influence of soil organic carbon fractions on the soil priming effect under different vegetation restoration modes

AbstractThe soil priming effect is a key mechanism influencing carbon (C) cycling processes between the forest soil organic carbon (SOC) pool and the atmosphere. Different vegetation restoration modes have different SOC pool compositions, and it is not clear whether such differences affect the soil priming effect. Therefore, we selected soil from six typical vegetation restoration modes Platycladus orientalis (PO), Pinus sylvestris (PS), Quercus acutissima (QA), shrub (SH) and wasteland (WL) in the soil and rocky mountainous areas of northern China and measured the soil properties, microbial communities, microbial necromass carbon (MNC), SOC fractions and the soil priming effect. The SOC content decreased in the order of PO (21.33 g kg−1), PS (22.00 g kg−1), QA (13.67 g kg−1), SH (13.33 g kg−1) and WL (10.33 g kg−1), and the trends of the mineral‐associated organic carbon (MAOC) and fungal necromass carbon (FNC) content were the same as those of the SOC content. The soil priming effect was greater in both forests and shrublands than in wastelands, with the greatest effect occurring in PO forests, where the soil priming effect reached 159.91 mg CO2‐C kg−1 soil after 30 days of incubation, which was 1.4 times greater than that in WL. The soil priming effect was mainly determined by the difference in MAOC content. In addition, the soil C/N ratio and bacterial community diversity also indirectly affected the soil priming effect by influencing the soil MNC and SOC fractions. Overall, afforestation increased the SOC content, fungal necromass contribution and mineral conservation, increasing the soil priming effect. This theoretical foundation supports the enhancement of SOC sequestration capacity by implementing various modes of vegetation restoration in the future.

Contribution of soil microbial necromass carbon to soil organic carbon in grassland under precipitation change and its influencing factors in loess hilly region, Northwest China.

To investigate the contribution of microbial necromass carbon (MNC) to soil organic carbon (SOC) and its influencing factors under precipitation changes in grassland, we conducted a precipitation experiment with seven different precipitation levels in the Bothriochloa ischaemum restoration area in the loess hilly region. We analyzed the contents and characteristics of fungal necromass carbon (FNC), bacterial necromass carbon (BNC), and MNC in different fractions of SOC under different treatments, including natural precipitation (CK), and increased and decreased 20%, 40%, 60% of natural precipitation (I20, I40, I60, D20, D40, D60) . The results showed that 1) MNC content in mineral organic carbon (MAOC) ranged from 1.62 g·kg-1 to 2.17 g·kg-1, which was higher than that in particulate organic carbon (POC) ranging from 0.69 g·kg-1 to 1.31 g·kg-1. The former was approximately 1.4 to 2.8 times as that of the latter. 2) FNC and MNC exhibited similar changes in both MAOC and POC fractions. BNC content in MAOC was approximately 1-3.1 times as that of FNC. FNC content in POC was generally higher than BNC except for I40 and I60 where BNC exceeded FNC. 3) Overall, both increases and decreases in precipitation resulted in elevated MNC/MAOC and BNC/MAOC ratios, but decreased MNC/POC and FNC/POC ratios. The MNC/MAOC ratios in I60 and D60 were 33.2% and 18.1% higher than CK, respectively. The BNC/MAOC ratios in D60, I40 and I60 were 28.0%, 23.0% and 19.1% higher than those in CK, respectively. Except for D60, the FNC/POC and MNC/POC ratios were significantly lower than CK under other treatments. In terms of POC fractions, the MNC/POC ratios of D40, D20, I20, I40 and I60 were 28.4%, 23.3%, 28.8%, 23.3% and 32.2% lower than that of CK, respectively. The FNC/POC ratio of D40, D20, I20, I40 and I60 was found to be lower by 23.3%, 16.1%, 21.0%, 27.0% and 31.0% compared to that of CK, respectively. 4) NH4+-N and pH were the primary factors influencing the content of MNC in different carbon fractions under varying precipitation conditions. In summary, alterations in precipitation (either increase or decrease) enhanced the contribution of BNC-dominated MNC to MAOC, but reduced the contribution of FNC-dominated MNC to POC. This study was of significance for understanding the distribution of microbial necromass across different organic carbon fractions under precipitation alterations.

Half substitution of mineral N with fish protein hydrolysate enhancing microbial residue C and N storage and climate benefits under high straw residue return

The widespread utilization of straw return was a popular practice straw disposal for highly intensive agriculture in China, which has brought about some negative impacts such as less time for straw complete biodegradation, aggravation of greenhouse gas evolution, and lower efficient of carbon accumulation. It was urgent to find an eco-friendly N-rich organic fertilizer instead of mineral N as activator to solve the above problems and lead a carbon accumulation in long tern management. Besides, microbial necromass was considered as a crucial contributor to persistent soil carbon (C) and nitrogen (N) pool. How organic fertilizer activators influence microbial residue under different amount of crop residues input remained unclear. Thus, soils incorporating moderate and high rate of rice straw residue with additions of half and full of organic activators (fish protein hydrolysates vs. manure) were incubated for measuring carbon dioxide (CO2) and nitrous oxide (N2O) emission, microbial community and necromass. It was found that soil CO2 emission was rapidest during the first 13 days of straw decomposition but remained lowest in the treatments of 50% mineral N substituted by fish protein hydrolysate. There were that 81%–89% of total CO2 release and 59%–65% of total N2O emission occurred within 60 days of incubation period, and bacterial community and nitrate positively affected soil CO2 and N2O release respectively. Straw incorporation amount and organic activator application interactively influenced soil CO2 emission but not affected soil N2O emission. After 360 days of incubation, the difference of bacterial necromass was noticeable but fungal necromass remained almost unaltered across all treatments. All treatments showed generally comparable contribution of microbial necromass N to the total N pool. The treatment of 50% mineral N substituted by fish protein hydrolysate under high rate of straw input (HSF50) promoted the highest proportion of microbial necromass C in soil organic C because of alleviating N limitation for microorganisms. Finally, HSF50 was recommended as an eco-friendly strategy for enhancing microbial necromass C and N storage and climate benefits in agroecosystems.

Transcriptomic analyses of bacterial growth on fungal necromass reveal different microbial community niches during degradation.

Bacteria are major drivers of organic matter decomposition and play crucial roles in global nutrient cycling. Although the degradation of dead fungal biomass (necromass) is increasingly recognized as an important contributor to soil carbon (C) and nitrogen (N) cycling, the genes and metabolic pathways involved in necromass degradation are less characterized. In particular, how bacteria degrade necromass containing different quantities of melanin, which largely control rates of necromass decomposition in situ, is largely unknown. To address this gap, we conducted a multi-timepoint transcriptomic analysis using three Gram-negative, bacterial species grown on low or high melanin necromass of Hyaloscypha bicolor. The bacterial species, Cellvibrio japonicus, Chitinophaga pinensis, and Serratia marcescens, belong to genera known to degrade necromass in situ. We found that while bacterial growth was consistently higher on low than high melanin necromass, the CAZyme-encoding gene expression response of the three species was similar between the two necromass types. Interestingly, this trend was not shared for genes encoding nitrogen utilization, which varied in C. pinensis and S. marcescens during growth on high vs low melanin necromass. Additionally, this study tested the metabolic capabilities of these bacterial species to grow on a diversity of C and N sources and found that the three bacteria have substantially different utilization patterns. Collectively, our data suggest that as necromass changes chemically over the course of degradation, certain bacterial species are favored based on their differential metabolic capacities.IMPORTANCEFungal necromass is a major component of the carbon (C) in soils as well as an important source of nitrogen (N) for plant and microbial growth. Bacteria associated with necromass represent a distinct subset of the soil microbiome and characterizing their functional capacities is the critical next step toward understanding how they influence necromass turnover. This is particularly important for necromass varying in melanin content, which has been observed to control the rate of necromass decomposition across a variety of ecosystems. Here we assessed the gene expression of three necromass-degrading bacteria grown on low or high melanin necromass and characterized their metabolic capacities to grow on different C and N substrates. These transcriptomic and metabolic studies provide the first steps toward assessing the physiological relevance of up-regulated CAZyme-encoding genes in necromass decomposition and provide foundational data for generating a predictive model of the molecular mechanisms underpinning necromass decomposition by soil bacteria.

Gradual and abrupt increase in atmospheric CO2 concentrations trigger divergent responses of microbial necromass accumulation in paddy soils

Microbial necromass C (MNC) plays a critical role in promoting soil organic C (SOC) formation and stabilization, particularly in the context of climate change. Most investigations on the impact of elevated CO2 concentrations (eCO2) on MNC have been performed by exposing ecosystems to an abrupt increase in CO2. However, the atmospheric CO2 increase is a gradual process, and knowledge about the response of necromass to gradual increase in CO2 is lacking. We tested the hypothesis that microbial necromass would show different responses to abrupt and gradual CO2 increases. Furthermore, it is still unknown whether eCO2 will trigger similar responses between surface and subsurface soil layers. Here, we determined the MNC concentrations and their proportions in SOC in surface and subsurface soil layers via open-top chambers. CO2 treatments included ambient control, abrupt CO2 increase by 200 ppm above control, and gradual CO2 increase by 40 ppm each year until reaching 200 ppm over five years. Overall, compared with the ambient control, abrupt eCO2 induced a significantly stronger decrease in necromass (20.3 %) when averaged across three soil layers, while gradual eCO2 led to insignificant variations in necromass (6.7 %). The necromass proportion in SOC decreased with depth under both eCO2 treatments with a significant decline occurring in abrupt eCO2 treatment. The observed greater magnitude of eCO2 effects on fungal necromass relative to bacterial necromass in subsurface layers illustrates a distinct microbial community response to climate change. Taken together, abrupt and gradual eCO2 approaches differed in their impact on MNC, and its response to eCO2 may be somewhat overestimated by abrupt eCO2 approach. Furthermore, MNC in subsurface soil witnessed a more sensitivity or vulnerability to eCO2 than that of topsoil. We suggest vigilant attention will need to be paid to the feedback of necromass C in deep soil poised by future climate change.

Variations in source-specific soil organic matter components across 32 forest sites in China

Forest soils store substantial amounts of carbon in various soil organic matter (SOM) components due to high plant litter inputs and active microbial turnover. However, the variations in plant- and microbial-derived SOM components in surface and subsurface forest soils across a wide geographic scale remain poorly understood. This study investigated the SOM components from aboveground and belowground plant inputs and fungal and bacterial necromass in surface (soil0–5 cm) and subsurface (soil5–10 cm) soils across 32 forest sites in China and analyzed their relationships with climate and edaphic factors. Compared to soil0–5 cm, soil5–10 cm exhibited lower soil organic carbon content and cutin biomarker concentration but higher concentrations of fungal necromass carbon and lignin phenols. Higher mean annual precipitation led to higher concentrations of cutin and suberin biomarkers in soil0–5 cm and soil5–10 cm, respectively. Higher soil organic carbon content was associated with lower plant-derived lignin biomarkers, higher lignin oxidation degrees, and increased microbial necromass-derived amino sugars across sites, highlighting the pivotal role of microbial necromass in SOM stabilization. Additionally, both fungal and bacterial necromass decreased with increasing mineral weathering across sites. These insights improve the understanding of environmental drivers of source-specific carbon storage in forest soils.Graphical

Variation of microbial necromass carbon and its potential relationship with humification during composting of chicken manure with and without biochar addition

Microbial necromass carbon (MNC) is an important stable organic C component. However, the variation of MNC and its potential relationship with humus components in composting remains uncertain. During a 45-day chicken manure composting study with and without biochar, MNC ranged from 24.9 to 77.9 g/kg and increased significantly by 80.9 % to 133 %. MNC constituted 5.77 % to 21.3 % of total organic C, with bacterial/fungal necromass C ratio ranging from 0.82 to 1.78. The MNC/humus C ratio ranged from 0.15 to 0.55, and humic acid C showed significant positive associations with bacterial necromass C (R2 = 0.72) and fungal necromass C (R2 = 0.51). Biochar addition reduced electrical conductivity and moisture content, increased pH, and induced microbial phosphorus limitation, thereby enhancing MNC content by 29.2 % and promoting humification. Our study is the first to elucidate the relationship between microbial necromass and humus substances, providing fundamental data for advancing the microbial carbon pump theory in composting.

Microbial life-history strategies and particulate organic carbon mediate formation of microbial necromass carbon and stabilization in response to biochar addition

Microbial necromass carbon (MNC) contributes significantly to the formation of soil organic carbon (SOC). However, the microbial carbon sequestration effect of biochar is often underestimated and influenced by nutrient availability. The mechanisms associated with the formation and stabilization of MNC remain unclear, especially under the combined application of biochar and nitrogen (N) fertilizer. Thus, in a long-term field experiment (11 years) based on biochar application, we utilized bacterial 16S rRNA gene sequencing, fungal ITS amplicon sequencing, metagenomics, and microbial biomarkers to examine the interactions between MNC accumulation and microbial metabolic strategies under combined treatment with biochar and N fertilizer. We aimed to identify the critical microbial modules and species involved, and to analyze the sites where MNC was immobilized from various components. Biochar application increased the MNC content by 13.9 %. Among the MNC components, fungal necromass contributed more to MNC, but bacteria were more readily enriched after biochar application. The microbial life-history strategies that affected MNC formation under the application of various amounts biochar were linked to the N application level. Under N added at 226.5 kg ha−1, communities such as Actinobacteria and Bacteroidetes with high-growth yield strategies were prevalent and contributed to MNC production. By contrast, under N added at 113.25 kg ha−1 with high biochar application, Proteobacteria with strong resource acquisition strategies were dominant and MNC accumulation was lower. The mineral-associated organic carbon pool was rapidly saturated with the addition of biochar, so the contribution of fungal necromass carbon may have been reduced by reutilization, thereby resulting in the more rapid preservation of bacterial necromass carbon in the particulate organic carbon pool. Overall, our findings indicate that microbial life history traits are crucial for linking microbial metabolic processes to the accumulation and stabilization of MNC, thereby highlighting the their importance for SOC accumulation in farmland soils, and the need to tailor appropriate biochar and N fertilizer application strategies for agricultural soils.

Fungal necromass contribution to carbon sequestration in global croplands: A meta-analysis of driving factors and conservation practices

Fungal necromass carbon (FNC) contributes significantly to the build-up of soil organic carbon (SOC) by supplying abundant recalcitrant polymeric melanin present in the fungal cell wall. However, the influence of a wide range of conservation practices and associated factors on FNC accumulation and contribution to SOC in global croplands remains unexplored. Here, a meta-analysis was performed using 873 observations across three continents, together with structural equation modeling, to evaluate conservation practices and factors responsible for the enhancement of FNC and SOC. FNC content (8.39 g kg−1) of North American soils was highest compared to FNC content of Asian and European soils. The structural equation models showed a significant (p < 0.05) positive influence of microbial biomass carbon (MBC), soil pH, and clay contents on the accumulation of FNC. Soil C/N ratio and climate factors, however, had only minor influences on FNC accumulation. Notably, the main driver of FNC was MBC, which is mainly influenced by the soil total N and geographic factors in the study areas. Typical 5 cropland practices had significant effect size (p < 0.05) on FNC, leading to an increase of 12 % to 26 %, and the FNC content was greatest under straw amendment (26 %). Fungal necromass accumulation efficiency ranged from 23 % to 45 % depending on cropland practices: non- and reduced tillage was the most efficient (45 %), followed by crop coverage (32 %), straw amendment (30 %), and manure application (27 %), while N fertilization had the lowest efficiency (23 %). We conclude that FNC contributes to over a quarter of SOC, highlighting its major role in enhancing C sequestration worldwide. Conservation practices, particularly non-tillage or reduced tillage, are important to enhance C sequestration from FNC in croplands.

Microbial necromass facilitated the humification process through amino sugar reactions during the co-composting of cow manure plus sawdust.

Humus (HS) reservoirs can embed microbial necromass (including cell wall components that are intact or with varying degrees of fragmentation) in small pores, raising widespread concerns about the potential for C/N interception and stability in composting systems. In this study, fresh cow manure and sawdust were used for microbial solid fermentation, and the significance of microbial residues in promoting humification was elucidated by measuring their physicochemical properties and analyzing their microbial informatics. These results showed that the stimulation of external carbon sources (NaHCO3) led to an increase in the accumulation of bacterial necromass C/N from 6.19 and 0.91µg/mg to 21.57 and 3.20µg/mg, respectively. Additionally, fungal necromass C/N values were about 3 times higher than the initial values. This contributed to the increase in HS content and the increased condensation of polysaccharides and nitrogen-containing compounds during maturation. The formation of cellular debris mainly depends on the enrichment of Actinobacteria, Proteobacteria, Ascomycota, and Chytridiomycota. Furthermore, Euryarchaeota was the core functional microorganism secreting cell wall lytic enzymes (including AA3, AA7, GH23, and GH15). In conclusion, this study comprehensively analyzed the transformation mechanisms of cellular residuals at different profile scales, providing new insights into C/N cycles and sequestration.

Differential carbon accumulation of microbial necromass and plant lignin by pollution of polyethylene and polylactic acid microplastics in soil

The wide microplastics (MPs) occurrence affects soil physicochemical and biological properties, thereby influencing its carbon cycling and storage. However, the regulation effect of MPs on soil organic carbon (SOC) formation and stabilization remains unclear, hindering the accurate prediction of carbon sequestration in future global changes under continuous MP pollution. Phospholipid fatty acids, amino sugars and lignin phenols were used in this study as biomarkers for microbial community composition, microbial necromass and plant lignin components, respectively, and their responses to conventional (polyethylene; PE) and biodegradable (polylactic acid; PLA) MPs were explored. Results showed PLA MPs had positive effects on soil microbial biomass, while the positive and negative effects of PE MPs on microbial biomass varied with MP concentration. PE and PLA MPs increased microbial necromass contents and their contribution to SOC, mainly due to the increase in fungal necromass. On the contrary, PE and PLA MPs reduced lignin phenols and their contribution to SOC, mainly owing to the reduction in vanillyl-type phenols. The response of microbial necromass to PLA MPs was higher than that to PE MPs, whereas the response of lignin phenols was the opposite. MPs increased SOC level, with 83%–200% and 50%–75% of additional SOC in PE and PLA treatments, respectively, originating from microbial necromass carbon. This finding indicates that the increase in SOC pool in the presence of MPs can be attributed to soil microbial necromass carbon, and MPs increased capacity and efficacy of microbial carbon pump by increasing microbial turnover and reducing microbial N limitation. Moreover, the increase in amino sugars to lignin phenols ratio in PE treatment was higher than that in PLA treatment, and the increase in SOC content in PLA treatment was higher than that in PE treatment, indicating a high possibility of SOC storage owing to PLA MPs.

Minimal impact of nitrogen addition on bacterial and fungal communities during fungal necromass decomposition in a subalpine coniferous plantation

Fungal necromass is a vital source of stable carbon and available nitrogen in soils, especially in coniferous forests that are extensively colonized by ectomycorrhizal fungi. Here, we examined changes in the necromass microbial community during a 20-week incubation experiment in a subalpine coniferous plantation. We also explored how the necromass microbial community responded to simulated nitrogen deposition, a global change factor that substantially affects nitrogen-limited coniferous forests. The results showed that necromass microbial community was more dominated by copiotrophic bacteria with high community average rRNA operon copy number (3.1–6.4) and fast-growing fungi (moulds and yeasts) than surrounding soils. Ectomycorrhizal fungi also constituted a significant portion of the fungal community during the fast decay stage (first 5 weeks) of necromass decomposition, indicating that fungal necromass may serve as a nitrogen source for the nitrogen-limited trees. The shifts in microbial functional compositions (e.g., decreased proportions of copiotrophic bacteria) as decomposition progressed were partly due to alterations in substrate chemistry (i.e., the levels of aliphatic compounds including soluble carbohydrate, amorphous glucan polymers, and crystalline glucans and chitin). In contrast, nitrogen addition had limited influences on microbial diversity and composition, likely due to the nitrogen-rich nature of the fungal necromass. Overall, our findings reinforce the existence of a specific and dynamic core necrobiome in fungal necromass and enhance our understanding of how fungal necrobiome responds to environmental changes.

Divergent accumulation of microbial necromass and plant lignin phenol induced by adding maize straw to fertilized soils

Plant carbon (C) inputs and their subsequent microbial transformation affect the build-up process of soil organic C (SOC) pool. Nevertheless, there are knowledge gaps on how crop straw addition modifies SOC composition at molecular-level, especially in soils with different fertilization practices. Here, long-term fertilized Mollisols (unfertilized control, NF; inorganic fertilization, IF; inorganic fertilization plus manure, IFM) were incubated with or without maize straw addition in a 900-day field mesocosm experiment. The microbial necromass and plant lignin components contents were synchronously quantified based on amino sugar and lignin phenol biomarkers, respectively. And changes in SOC chemical composition were examined using solid-state 13C nuclear magnetic resonance spectroscopy. Relative to the NF treatment, long-term fertilization increased amino sugar and lignin phenol contents, and manure application enhanced the accumulation of plant lignin components more than that of microbial necromass. Compared with the NF treatment, the IF treatment decreased the relative proportion of alkyl C and SOC content, suggesting that changes in microbial necromass and lignin phenol were not always consistent with changes in SOC. For the NF and IF treatments, maize straw incorporation increased both amino sugar content and its contribution to SOC, indicating that microbial anabolism was important for SOC accumulation after adding maize straw in C-poor soils. For the IFM treatment, maize straw addition decreased lignin phenol content (27 %) and its contribution to SOC but increased the contribution of amino sugar to SOC, reflecting an enlarged contribution of microbial necromass to soil C pool formation under manure application after maize straw addition. The contribution of lignin phenol to SOC was decreased from days 360–900, whereas fungal necromass C content and the contribution of amino sugar to SOC were increased from days 360–900 under the IFM soil with maize straw addition, indicating that plant lignin components might be converted into fungal biomass and its necromass with increasing maize straw decomposition under manure application. Overall, we concluded that maize straw addition favored microbial immobilization in all fertilization treatments.

Fungi rather than bacteria drive early mass loss from fungal necromass regardless of particle size.

Microbial necromass is increasingly recognized as an important fast-cycling component of the long-term carbon present in soils. To better understand how fungi and bacteria individually contribute to the decomposition of fungal necromass, three particle sizes (>500, 250-500, and <250 μm) of Hyaloscypha bicolor necromass were incubated in laboratory microcosms inoculated with individual strains of two fungi and two bacteria. Decomposition was assessed after 15 and 28 days via necromass loss, microbial respiration, and changes in necromass pH, water content, and chemistry. To examine how fungal-bacterial interactions impact microbial growth on necromass, single and paired cultures of bacteria and fungi were grown in microplates containing necromass-infused media. Microbial growth was measured after 5 days through quantitative PCR. Regardless of particle size, necromass colonized by fungi had higher mass loss and respiration than both bacteria and uninoculated controls. Fungal colonization increased necromass pH, water content, and altered chemistry, while necromass colonized by bacteria remained mostly unaltered. Bacteria grew significantly more when co-cultured with a fungus, while fungal growth was not significantly affected by bacteria. Collectively, our results suggest that fungi act as key early decomposers of fungal necromass and that bacteria may require the presence of fungi to actively participate in necromass decomposition.

Long-term residue returning increased subsoil carbon quality in a rice-wheat cropping system

Residue returning (RR) was widely implemented to increase soil organic carbon (SOC) in farmland. Extensive studies concentrated on the effects of RR on SOC quantity instead of SOC fractions at aggregate scales. This study investigated the effects of 20-year RR on the distribution of labile (e.g., dissolved, microbial biomass, and permanganate oxidizable organic) and stable (e.g., microbial necromass) carbon fractions at aggregate scales, as well as their contribution to SOC accumulation and mineralization. The findings indicated a synchronized variation in the carbon content of bacterial and fungal necromass. Residue retention (RR) notably elevated the concentration of bacterial and fungal necromass carbon, while it did not amplify the microbial necromass carbon (MNC) contribution to SOC when compared to residue removal (R0) in the topsoil (0–5 cm). In the subsoil (5–15 cm), RR increased the MNC contribution to SOC concentration by 21.2%–33.4% and mitigated SOC mineralization by 12.6% in micro-aggregates (P < 0.05). Besides, RR increased soil β-glucosidase and peroxidase activities but decreased soil phenol oxidase activity in micro-aggregates (P < 0.05). These indicated that RR might accelerate cellulose degradation and conversion to stable microbial necromass C, and thus RR improved SOC stability because SOC occluded in micro-aggregates were more stable. Interestingly, SOC concentration was mainly regulated by MNC, while SOC mineralization was by dissolved organic carbon under RR, both of which were affected by soil carbon, nitrogen, and phosphorus associated nutrients and enzyme activities. The findings of this study emphasize that the paths of RR-induced SOC accumulation and mineralization were different, and depended on stable and labile C, respectively. Overall, long-term RR increased topsoil carbon quantity and subsoil carbon quality.

Long‐term successive biochar application increases plant lignin and microbial necromass accumulation but decreases their contributions to soil organic carbon in rice–wheat cropping system

AbstractBiochar application is widely recognized as an effective approach for increasing soil organic carbon (SOC) and mitigating climate change in agroecosystems. However, the effects of biochar application on net accumulations and relative contributions of different SOC sources remain unclear. Here, we explored the effects of biochar application on plant‐derived (PDC) and microbial necromass C (MNC) in a 10‐year experimental rice–wheat rotation field receiving four different intensities of biochar application (0, 2.25, 11.5, and 22.5 t ha−1 for each crop season), using phospholipid fatty acids (PLFAs), lignin phenols and amino sugars as biomarkers of microbial biomass, PDC and MNC, respectively. Our results showed that biochar application increased SOC content and stock by 32.6%–203% and 26.4%–145%, respectively. Higher biochar application (11.5 and 22.5 t ha−1) increased soil pH, total nitrogen (TN), total phosphorus (TP), SOC/TN, and root biomass. In addition, higher biochar application enhanced bacterial, fungal, and total microbial biomass. Plant lignin phenols and MNC contents significantly increased, whereas their contributions to SOC significantly decreased with the increase in biochar application rates due to the disproportionate increase in PDC and MNC, and SOC. Fungal necromass had a greater contribution to SOC than bacterial necromass. The fungal/bacterial necromass decreased from 2.56 to 2.26 with increasing biochar application rates, because of the higher abundances of bacteria than that of fungi as indicated by PLFAs under higher biochar application rates. Random forest analyses revealed that pH, TP, and SOC/TN were the main factors controlling plant lignin and MNC accumulation. Structural equation modeling revealed that biochar application increased lignin phenols by stimulating root biomass, whereas enhanced MNC accumulation was primarily from increased microbial biomass and lignin phenols. Overall, our findings suggest that biochar application increases the accumulation of the two SOC sources but decreases their contributions to SOC in paddy soils.