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.

Long-term soil warming decreases soil microbial necromass carbon by adversely affecting its production and decomposition.

Microbial necromass carbon (MNC) accounts for a large fraction of soil organic carbon (SOC) in terrestrial ecosystems. Yet our understanding of the fate of this large carbon pool under long-term warming is uncertain. Here, we show that 14 years of soil warming (+4°C) in a temperate forest resulted in a reduction in MNC by 11% (0-10 cm) and 33% (10-20 cm). Warming caused a decrease in the content of MNC due to a decline in microbial biomass carbon and reduced microbial carbon use efficiency. This reduction was primarily caused by warming-induced limitations in available soil phosphorus, which, in turn, constrained the production of microbial biomass. Conversely, warming increased the activity of soil extracellular enzymes, specifically N-acetylglucosaminidase and leucine aminopeptidase, which accelerated the decomposition of MNC. These findings collectively demonstrate that decoupling of MNC formation and decomposition underlie the observed MNC loss under climate warming, which could affect SOC content in temperate forest ecosystems more widespread.

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 ß-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.

Salinity decreases the contribution of microbial necromass to soil organic carbon pool in arid regions.

Saline soils are widely distributed in arid areas but there is a lack of mechanistic understanding on the effect of salinity on the formation and biochemical composition of soil organic carbon (SOC). We investigated the effects of salinity on the accumulation of microbial necromass under natural vegetation and in cropland in salt-affected arid areas stretching over a 1200-km transect in northwest China. Under both natural vegetation and cropland, microbial physiological activity (indicated by microbial biomass carbon normalized enzymatic activity) decreased sharply where the electrical conductivity approached 4 ds m-1 (a threshold to distinguish between saline and non-saline soils), but microbial biomass was only slightly affected by salinity. These indicated that a larger proportion of microbes could be inactive or dormant in saline soils. The contribution of fungal necromass C to SOC decreased but the contribution of bacterial necromass C to the SOC increased with increasing soil salinity. Adding fungal and bacterial necromass C together, the contribution of microbial necromass C to SOC in saline soils was 32-39 % smaller compared with non-saline soils. Fungal necromass C took up 85-86 % of microbial necromass C in non-saline soils but this proportion dropped to 60-66 % in saline soils. We suggested that the activity, growth, and turnover rate of microbes slowed by salinity was responsible for the decreased accumulation of fungal necromass in saline compared with non-saline soils, while the increased accumulation of bacterial residue in saline soils could be induced mainly by its slower decomposition. Soil microbial biomass was a poor predictor for the accumulation of microbial necromass in saline soils. We demonstrated a reduced contribution of microbial necromass to SOC and a shift in its composition towards the increase in bacterial origin in saline relative to non-saline soils. We concluded that salinity profoundly changes the biochemistry of SOC in arid regions.

[Response of soil microbial necromass carbon to litter and root carbon inputs in a mid-subtropical Castanopsis carlesii plantation]. / ä¸­äºšçƒ­å¸¦ç±³æ§ äººå·¥æž—åœŸå£¤å¾®ç”Ÿç‰©æ®‹ä½“ç¢³å¯¹å‡‹è½ç‰©å’Œæ ¹ç³»ç¢³è¾“å ¥çš„å“åº”.

Microbial necromass carbon (MNC) is a crucial source for stable soil carbon pool, and understanding its response to carbon inputs from both aboveground (litter) and belowground (roots) in subtropical forest soils is essential for assessing soil carbon stocks in global ecosystems. In a Castanopsis carlesii plantation at the Sanming Forest Ecosystem National Observation and Research Station in Fujian Province, we conducted an experiment with five treatments, including root removal (NR), aboveground litter removal (NL), no litter input (removals of both aboveground litter and roots, NI), double aboveground litter addition (DL), and control (CK). After seven years, we collected soil samples in the 0-10 cm soil layer to examine changes in MNC content and its contribution to soil organic carbon (SOC). Results showed that NR treatment reduced MNC, bacterial necromass carbon (BNC), and fungal necromass carbon (FNC) by 15.9%, 20.2%, and 14.5%, respectively, while other treatments did not induce significant changes. The NR, NL, NI, and DL treatments did not affect the contributions of BNC, FNC, and MNC to SOC. Correlation and path analyses revealed that litter and root carbon input treatments could alter the MNC content directly or indirectly through changing soil available substrates and microbial community structure. Our results suggested that roots exert a stronger influence on the maintenance of MNC than aboveground carbon source in the mid-subtropical plantations.

[Effects of mixed broadleaved tree species with pure Pinus massoniana plantation on soil microbial necromass carbon and organic carbon fractions]. / é©¬å°¾æ¾çº¯æž—æ··äº¤æ”¹é€ å¯¹åœŸå£¤å¾®ç”Ÿç‰©æ®‹ä½“ç¢³å’Œæœ‰æœºç¢³ç»„åˆ†çš„å½±å“.

Mixing native broadleaved tree species is a widely used method for renovating Pinus massoniana plantations. Soil microbial necromass carbon and organic carbon fractions are important parameters for evaluating the impacts of tree species mixing and soil organic carbon (SOC) stability. However, their responses to the mixing and renovation of P. massoniana plantation has not been understood yet. Here, we selected a pure P. massoniana plantation (PP) and a mixed P. massoniana and Castanopsis hystrix plantation, with ages of 16 (MP16) and 38 years (MP38), respectively, as the research objects. We quantified soil physical and chemical properties, microbial necromass carbon content, and organic carbon components at different soil layers to reveal whether and how the introduction of C. hystrix into P. massoniana plantation affected soil microbial necromass carbon and organic carbon components. The results showed that the mixed P. massoniana and C. hystrix plantation significantly reduced fungal necromass carbon content and the ratio of fungal/bacterial necromass carbon in the 0-20 cm and 20-40 cm soil layers. There were no significant differences in microbial necromass carbon contents, bacterial necromass carbon contents, and their contributions to SOC among the different plantations. The contribution of fungal necromass carbon to SOC was higher than that of bacterial necromass carbon in all plantation types. The contribution of soil mineral-associated organic carbon (MAOC) to SOC was higher than that of occluded particulate organic carbon (oPOC) and light-free particulate organic carbon (fPOC) for all plantation types. Mixing the precious broadleaved tree species (i.e., C. hystrix) with coniferous species (P. massoniana) significantly increased MAOC content and the contribution of MAOC, oPOC, and fPOC to SOC in the 0-20 cm and 20-40 cm soil layers. The MAOC of MP38 was significantly higher than that of PP in all soil layers and the MAOC of MP38 stands were significantly higher than MP16 stands in the 20-40 cm, 40-60 cm, and 60-100 cm soil layers, indicating that hybridization enhanced SOC stability and that the SOC of MP38 stands were more stable than MP16 stands. SOC and total nitrogen contents were the main environmental factors driving the changes in soil microbial necromass carbon, while soil total nitrogen and organically complexed Fe-Al oxides were the primary factors affecting organic carbon fraction. Therefore, SOC stability can be enhanced by introducing native broadleaved species, such as C. hystrix, during the management of the P. massoniana plantation.

Spatial variation and controls of soil microbial necromass carbon in a tropical montane rainforest.

Soil microbial necromass carbon is an important component of the soil organic carbon (SOC) pool which helps to improve soil fertility and texture. However, the spatial pattern and variation mechanisms of fungal- and bacterial-derived necromass carbon at local scales in tropical rainforests are uncertain. This study showed that microbial necromass carbon and its proportion in SOC in tropical montane rainforest exhibited large spatial variation and significant autocorrelation, with significant high-high and low-low clustering patterns. Microbial necromass carbon accounted for approximately one-third of SOC, and the fungal-derived microbial necromass carbon and its proportion in SOC were, on average, approximately five times greater than those of bacterial-derived necromass. Structural equation models indicated that soil properties (SOC, total nitrogen, total phosphorus) and topographic features (elevation, convexity, and aspect) had significant positive effects on microbial necromass carbon concentrations, but negative effects on its proportions in SOC (especially the carbon:nitrogen ratio). Plant biomass also had significant negative effects on the proportion of microbial necromass carbon in SOC, but was not correlated with its concentration. The different spatial variation mechanisms of microbial necromass carbon and their proportions in SOC are possibly related to a slower accumulation rate of microbial necromass carbon than of plant-derived organic carbon. Geographic spatial correlations can significantly improve the microbial necromass carbon model fit, and low sampling resolution may lead to large uncertainties in estimating soil carbon dynamics at specific sites. Our work will be valuable for understanding microbial necromass carbon variation in tropical forests and soil carbon prediction model construction with microbial participation.

Cross-scale spatial variability and associations of carbon pools provide insight into regulating carbon sequestration in tropical montane rainforests.

The spatial distribution of plant, soil, and microbial carbon pools, along with their intricate interactions, presents a great challenge for the current carbon cycle research. However, it is not clear what are the characteristics of the spatial variability of these carbon pools, particularly their cross-scale relationships. We investigated the cross-scale spatial variability of microbial necromass carbon (MNC), soil organic carbon (SOC) and plant biomass (PB), as well as their correlation in a tropical montane rainforest using multifractal analysis. The results showed multifractal spatial variations of MNC, SOC, and PB, demonstrating their adherence to power-law scaling. MNC, especially low MNC, exhibited stronger spatial heterogeneity and weaker evenness compared with SOC and PB. The cross-scale correlation between MNC and SOC was stronger than their correlations at the measurement scale. Furthermore, the cross-scale spatial variability of MNC and SOC exhibited stronger and more stable correlations than those with PB. Additionally, this research suggests that when SOC and PB are both low, it is advisable for reforestations to potentiate MNC formation, whereas when both SOC and PB are high some thinning can be advisable to favour MNC formation. Thus, these results support the utilization of management measures such as reforestation or thinning as nature-based solutions to regulate carbon sequestration capacity of tropical forests by affecting the correlations among various carbon pools.

Contribution of microbial necromass to soil organic carbon formation during litter decomposition under incubation conditions.

We conducted a 512-day incubation experiment to study the dynamics of microbial necromass and soil carbon fraction in the 'litter-soil' transformation interface soil layer (TIS) during litter decomposition, using a perennial C3 herb, Stipa bungeana, in the loess hills. The results showed that soil microbial necromass was dominated by fungi in the early and middle stages, and by bacteria in the late stage. The contribution of fungal necromass C to mineral-associated organic C (MAOC) was significantly higher (38.7%-75.8%) than that of bacteria (9.2%-22.5%) and 2-3 times more than the contribution rate of bacterial necromass. Soil organic C (SOC) content was decreasing during litter decomposition. The input of plant C resources stimulated microbial utilization of soil C fractions. The continuous decrease in particulate organic C during the early and late stages of decomposition was directly responsible for the decrease in SOC content. In contrast, the fluctuating changes in microbial necromass C and MAOC played an indirect role in the reduction of SOC. The increase in soil microbial necromass C caused by a single exogenous addition of litter did not directly contribute to SOC accumulation.

[Impacts of Land Use Intensification Level on Fluvo-aquic Cropland Soil Microbial Community Abundance and Necromass Accumulation in North China].

Despite the essential role of soil microorganisms in nutrient turnover in soil ecological systems and the recognized paramount significance of microbial necromass to soil organic carbon accumulation, how microbial community abundance and necromass respond to land use intensification level regulation remains poorly understood. To address this knowledge gap, based on the land use intensification level, three treatments were set up[annual wheat-maize rotation (CC), alternating temporary grassland with wheat planting (TG), and perennial grassland (PG)], and a long-term fixed filed experiment was established to investigate the influences of the regulation of land use intensification level on bacterial and fungal community abundances; the accumulation of bacterial, fungal, and total microbial necromass; and their contributions to SOC sequestration using droplet digital PCR and amino sugar detection technologies. We further sought to determine the key factors driving the bacterial, fungal, and total microbial necromass C accumulation. Our results demonstrated that fungal community abundance was strongly affected by land use intensification level regulation compared to that of the bacterial community, which increased with decreasing land use intensification level. The total microbial necromass C predominated the SOC accumulation across all three land use intensification levels, which contributed 52.78%, 58.36%, and 68.87% to SOC, respectively, exhibiting an increasing trend with the decline in land use intensification level. Fungal necromass C accounted for more than 80% of the total microbial necromass C, indicating its predominance in the accumulation of the total microbial necromass C and active variation via the reduction in land use intensification level. There was no significant difference in bacterial necromass C (MurA) content, with the trend of CC

Divergent control and variation in bacterial and fungal necromass carbon respond to the abandonment of rice terraces.

The abandonment of rice terraces in hilly agroecosystems in recent decades has caused substantial changes in microbial characteristics and their impact on microbial necromass carbon (MNC) and soil organic carbon (SOC). Nevertheless, the regulatory mechanisms and impact pathways of MNC remain unclear. Here, soil samples were collected from 0 to 120 cm soil profiles in rice terraces, dry land (DL), and forest land (FL) for analysis. After converting rice terraces to DL and FL, MNC decreased significantly by 31.12% and 38.33%, while SOC decreased significantly by 51.26% and 29.87% respectively. These reductions are due to the loss of terrace management practices and associated functions. There were no significant changes in bacterial necromass carbon (BNC), whereas fungal necromass carbon (FNC) experienced a significant decrease. As a result, the decline in SOC may be primarily attributed to the reduction in FNC. BNC and FNC were regulated by bacterial life history strategies and fungal biomass, respectively. However, bacterial copiotrophs experienced a significant reduction after rice terrace abandonment. The regulation of BNC may be influenced by other factors, potentially offsetting the negative impact of abandonment. Dissolved organic carbon and bulk density were the primary control factors for bacterial community composition and fungal biomass, respectively. Additionally, the impact of soil layers on the alterations in MNC and SOC was more significant compared to the abandonment of rice terraces. These findings indicate that short-term abandonment of rice terraces results in a decrease in SOC, potentially compromising the ecological service function of the hilly agroecosystems. In the face of rapid population growth and global warming, it is crucial to minimize terrace abandonment and enhance utilization rates. This approach will effectively support sustainable terrace management and ecological services.

The effect of thinning intensity on the soil carbon pool mediated by soil microbial communities and necromass carbon in coastal zone protected forests.

Thinning is a common forest management measure that can effectively maintain the ecological service function of protected forests. However, the effect of thinning on the soil carbon (C) pool remains uncertain. In particular, we lack an understanding of the complete link between thinning and microbial communities, microbial necromass C, and consequently, soil C pools in coastal zone protected forests. In this study, three thinning intensities, i.e., a control treatment (CT, i.e., no thinning), light thinning (LT) and heavy thinning (HT), were established in three types of forests (Quercus acutissima Carruth, Pinus thunbergii Parl and mixed Quercus acutissima Carruth and Pinus thunbergii Parl, i.e., QAC, PTP and QP, respectively). Two years after the completion of thinning, we investigated the changes in the soil organic carbon (SOC) fractions, soil microbial community and soil microbial necromass C in the surface layer (0-20 cm) and thoroughly evaluated the relationship between the potential change in SOC and the microbial community. Compared with CT, there was no change in the SOC content under LT and HT, but thinning conducted in QAC increased the proportion of mineral-associated organic C (MAOC) in SOC. Moreover, both LT and HT reduced the soil carbon lability (CL) in the QAC and QP forests. Different thinning intensities changed the soil microbial community structure, and most of the variation was explained by thinning and the soil physicochemical properties. The proportion of soil bacterial and fungal necromass C to SOC increased with increasing thinning intensity. The content of soil bacterial and fungal necromass C was mainly controlled by the relative abundance of the core phylum (relative abundance>10 %). Thinning affected the soil C pool by affecting the content of soil bacterial and fungal necromass C, but their accumulation pathways was different. The results showed that thinning was beneficial to the stability of SOC. The microbial C pool, total organic C pool and even bacterial and fungal C pools should be distinguished when studying the soil C pool, which can effectively deepen our understanding of the mechanism by which soil microorganisms affect the soil C pool.

Dramatic Carbon Loss in a Permafrost Thaw Slump in the Tibetan Plateau is Dominated by the Loss of Microbial Necromass Carbon.

Thaw slumps can lead to considerable carbon loss in permafrost regions, while the loss of components from two major origins, i.e., microbial and plant-derived carbon, during this process remains poorly understood. Here, we provide direct evidence that microbial necromass carbon is a major component of lost carbon in a retrogressive permafrost thaw slump by analyzing soil organic carbon (SOC), biomarkers (amino sugars and lignin phenols), and soil environmental variables in a typical permafrost thaw slump in the Tibetan Plateau. The retrogressive thaw slump led to a ∼61% decrease in SOC and a ∼25% SOC stock loss. As evident in the levels of amino sugars (average of 55.92 ± 18.79 mg g-1 of organic carbon, OC) and lignin phenols (average of 15.00 ± 8.05 mg g-1 OC), microbial-derived carbon (microbial necromass carbon) was the major component of the SOC loss, accounting for ∼54% of the SOC loss in the permafrost thaw slump. The variation of amino sugars was mainly related to the changes in soil moisture, pH, and plant input, while changes in lignin phenols were mainly related to the changes in soil moisture and soil bulk density.

Climate warming alters the relative importance of plant root and microbial community in regulating the accumulation of soil microbial necromass carbon in a Tibetan alpine meadow.

Climate warming is predicted to considerably affect variations in soil organic carbon (SOC), especially in alpine ecosystems. Microbial necromass carbon (MNC) is an important contributor to stable soil organic carbon pools. However, accumulation and persistence of soil MNC across a gradient of warming are still poorly understood. An 8-year field experiment with four levels of warming was conducted in a Tibetan meadow. We found that low-level (+0-1.5°C) warming mostly enhanced bacterial necromass carbon (BNC), fungal necromass carbon (FNC), and total MNC compared with control treatment across soil layers, while no significant effect was caused between high-level (+1.5-2.5°C) treatments and control treatments. The contributions of both MNC and BNC to soil organic carbon were not significantly affected by warming treatments across depths. Structural equation modeling analysis demonstrated that the effect of plant root traits on MNC persistence strengthened with warming intensity, while the influence of microbial community characteristics waned along strengthened warming. Overall, our study provides novel evidence that the major determinants of MNC production and stabilization may vary with warming magnitude in alpine meadows. This finding is critical for updating our knowledge on soil carbon storage in response to climate warming.

Responses of microbial necromass carbon and microbial community structure to straw- and straw-derived biochar in brown earth soil of Northeast China.

Soil microbial organisms are conducive to SOC sequestration. However, little attention has been given to the contributions of living MBC and microbial necromass carbon to the SOC pool under biochar and straw amendments. The aims of the study were to explore (1) the effects of maize straw and biochar on MBC, POC, MAOC, DOC and microbial necromass carbon; (2) the contribution of MBC and microbial necromass carbon to the SOC pool; and (3) the relationships among the soil microbial community structure, microbial necromass carbon and other SOC fractions under maize straw and biochar application for nine consecutive years. Three treatments were studied: CK (applied chemical fertilizer only), BC (biochar applied annually at a rate of 2.625 t ha-1 combined with chemical fertilizer), and SR (straw applied annually at a rate of 7.5 t ha-1). Both biochar and straw increased the SOC contents after nine successive maize plant seasons; the DOC and MAOC contents were also increased by biochar and straw amendments. Biochar had advantages in increasing POC contents compared to straw. Biochar and straw increased MBC contents by 48.54% and 60.83% compared to CK, respectively. Straw significantly increased the Galn, GluN, MurA, ManN and total amino contents (P < 0.05); however, biochar significantly increased the Galn and GluN contents (P < 0.05) but had no impact on the MurA contents and decreased the ManN contents. Biochar mainly increased the fungal-derived necromass carbon contents but had no effect on the bacterial-derived necromass carbon, and straw increased both the bacterial- and fungal-derived necromass carbon contents. Straw had no influence on the ratios of microbial necromass carbon accounting for SOC and MAOC, but biochar decreased the ratios in the current study. Similarly, biochar mainly increased the fungal PLFA and total PLFA contents compared to CK, but straw increased bacterial PLFAs, fungal PLFAs and Actinomycetes PLFAs. Maize yield were increased by 7.44 and 9.16% by biochar and straw application, respectively. These results indicate that biochar stimulates fungal activities and turnover to contribute to the stable soil carbon pool and that biochar also improves POC contents to improve the soil organic carbon sink.