Ectomycorrhizal fungi redirect soil carbon stabilization pathways under nitrogen enrichment by regulating microbial necromass formation

Wenyuan He , Yu Zhang , Hongjian Wei , Tingying Xu , Wentao Hu , Hui Chen , Ming Tang

Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) : 146

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Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) :146 DOI: 10.1007/s11676-026-02082-3
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Ectomycorrhizal fungi redirect soil carbon stabilization pathways under nitrogen enrichment by regulating microbial necromass formation
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Abstract

Atmospheric nitrogen (N) deposition is altering forest soil carbon dynamics, yet the role of ectomycorrhizal (ECM) symbiosis in regulating soil organic carbon (SOC) stabilization under elevated N remains poorly understood. We investigated how N addition (0, 5, 10 and 20 mM urea) and ECM colonization interact to influence SOC partitioning, microbial functioning, and stabilization mechanisms in Eucalyptus grandis ×  Eucalyptus urophylla soils. ECM colonization remained consistently high across the N gradient, but fungal biomass declined with increasing N, indicating shifts in fungal allocation strategies. N addition alone favored accumulation of transient particulate organic carbon, whereas ECM inoculation promoted mineral-associated organic carbon (MAOC) at intermediate N levels, despite reductions in SOC hydrophobicity and chemical recalcitrance, suggesting that ECM-driven stabilization operates independently of chemical persistence. ECM fungi constrained microbial oxidative enzyme investment, notably suppressing polyphenol oxidase, thereby reshaping microbial carbon processing. Plant-derived CAZymes dominated SOC depolymerization, while SOC partitioning was independently regulated by a BradyrhizobiumPenicillium-centered microbial guild and by the CAZy enzyme families GH104 (bacterial) and GH17 (fungal). MAOC accumulation was more closely associated with microbial necromass formation and microbial carbon pump efficiency than with lignin chemistry. Collectively, these findings demonstrate that ECM symbiosis mediates soil carbon fate under N enrichment by coordinating microbial community composition, enzymatic activity, and necromass pathways, redirecting carbon from rapid mineralization to biologically mediated stabilization. This study highlights ECM fungi as critical biotic controllers of SOC persistence in N-enriched plantation ecosystems.

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Keywords

Ectomycorrhizal symbiosis / Nitrogen enrichment / Soil carbon pools / Microbial community / CAZymes / Eucalyptus

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Wenyuan He, Yu Zhang, Hongjian Wei, Tingying Xu, Wentao Hu, Hui Chen, Ming Tang. Ectomycorrhizal fungi redirect soil carbon stabilization pathways under nitrogen enrichment by regulating microbial necromass formation. Journal of Forestry Research, 2026, 37 (1) : 146 DOI:10.1007/s11676-026-02082-3

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References

[1]

Allison SD, Vitousek PM. Responses of extracellular enzymes to simple and complex nutrient inputs. Soil Biol Biochem, 2005, 37(5): 937-944.

[2]

Averill C, Hawkes CV. Ectomycorrhizal fungi slow soil carbon cycling. Ecol Lett, 2016, 19(8): 937-947.

[3]

Averill C, Turner BL, Finzi AC. Mycorrhiza-mediated competition between plants and decomposers drives soil carbon storage. Nature, 2014, 505(7484): 543-545.

[4]

Bååth E, Nilsson LO, Göransson H, Wallander H. Can the extent of degradation of soil fungal mycelium during soil incubation be used to estimate ectomycorrhizal biomass in soil?. Soil Biol Biochem, 2004, 36(12): 2105-2109.

[5]

Bera T, Vardanyan L, Inglett KS, Reddy KR, O’Connor GA, Erickson JE, Wilkie AC. Influence of select bioenergy by-products on soil carbon and microbial activity: a laboratory study. Sci Total Environ, 2019, 653: 1354-1363.

[6]

Bödeker ITM, Clemmensen KE, de Boer W, Martin F, Olson Å, Lindahl BD. Ectomycorrhizal Cortinarius species participate in enzymatic oxidation of humus in northern forest ecosystems. New Phytol, 2014, 203(1): 245-256.

[7]

Buchfink B, Xie C, Huson DH. Fast and sensitive protein alignment using DIAMOND. Nat Methods, 2015, 12(1): 59-60.

[8]

Cantarel BL, Coutinho PM, Rancurel C, Bernard T, Lombard V, Henrissat B. The carbohydrate-active EnZymes database (CAZy): an expert resource for glycogenomics. Nucleic Acids Res, 2009, 37: D233-D238.

[9]

Chen J, Elsgaard L, van Groenigen KJ, Olesen JE, Liang Z, Jiang Y, Lærke PE, Zhang YF, Luo YQ, Hungate BA, Sinsabaugh RL, Jørgensen U. Soil carbon loss with warming: new evidence from carbon-degrading enzymes. Glob Change Biol, 2020, 26(4): 1944-1952.

[10]

Chen SG, Zhang YQ, Ma J, Bai MY, Long JX, Liu M, Chen YL, Guo JB, Chen L. Contribution of soil microbial necromass carbon to soil organic carbon fractions and its influencing factors in different grassland types. Soil, 2025, 11(2): 883-898.

[11]

Chen X, Cao JJ, Sinsabaugh RL, Moorhead DL, Bardgett RD, Fanin N, Nottingham AT, Zheng XH, Chen J. Soil extracellular enzymes as drivers of soil carbon storage under nitrogen addition. Biol Rev, 2025, 100(4): 1716-1733.

[12]

Cheng YD, Huang JJ, Wang SL, Xiong K, Liang K, Wang FC, Wang SN, Zhang HP, Wang GG, Chen FS. Exploring the boost by dominant ectomycorrhizal trees to soil organic carbon sequestration in the subtropical forest of the Jiulianshan National Nature Reserve. J for Res, 2025, 36(1): 131.

[13]

Choreño-Parra EM, Treseder KK. Mycorrhizal fungi modify decomposition: a meta-analysis. New Phytol, 2024, 242(6): 2763-2774.

[14]

Clemmensen KE, Bahr A, Ovaskainen O, Dahlberg A, Ekblad A, Wallander H, Stenlid J, Finlay RD, Wardle DA, Lindahl BD. Roots and associated fungi drive long-term carbon sequestration in boreal forest. Science, 2013, 339(6127): 1615-1618.

[15]

Cotillard A, Kennedy SP, Kong LC, Prifti E, Pons N, Le Chatelier E, Almeida M, Quinquis B, Levenez F, Galleron N, Gougis S, Rizkalla S, Batto JM, Renault P, Doré J, Zucker JD, Clément K, Ehrlich SD. Dietary intervention impact on gut microbial gene richness. Nature, 2013, 500(7464): 585-588.

[16]

Cotrufo MF, Wallenstein MD, Boot CM, Denef K, Paul E. The Microbial Efficiency-Matrix Stabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: do labile plant inputs form stable soil organic matter?. Glob Change Biol, 2013, 19(4): 988-995.

[17]

Cotrufo MF, Soong JL, Horton AJ, Campbell EE, Haddix ML, Wall DH, Parton WJ. Formation of soil organic matter via biochemical and physical pathways of litter mass loss. Nat Geosci, 2015, 8(10): 776-779.

[18]

Dai ZM, Zang HD, Chen J, Fu YY, Wang XH, Liu HT, Shen CC, Wang JJ, Kuzyakov Y, Becker JN, Hemp A, Barberán A, Gunina A, Chen HH, Luo Y, Xu JM. Metagenomic insights into soil microbial communities involved in carbon cycling along an elevation climosequences. Environ Microbiol, 2021, 23(8): 4631-4645.

[19]

Domeignoz-Horta LA, Shinfuku M, Junier P, Poirier S, Verrecchia E, Sebag D, DeAngelis KM. Direct evidence for the role of microbial community composition in the formation of soil organic matter composition and persistence. ISME Commun, 2021, 1(1): 64.

[20]

Fang YY, Nazaries L, Singh BK, Singh BP. Microbial mechanisms of carbon priming effects revealed during the interaction of crop residue and nutrient inputs in contrasting soils. Glob Change Biol, 2018, 24(7): 2775-2790.

[21]

Feng Q, Liang SS, Jia HJ, Stadlmayr A, Tang LQ, Lan Z, Zhang DY, Xia HH, Xu XY, Jie ZY, Su LL, Li XP, Li X, Li JH, Xiao L, Huber-Schönauer U, Niederseer D, Xu X, Al-Aama JY, Yang HM, Wang J, Kristiansen K, Arumugam M, Tilg H, Datz C, Wang J. Gut microbiome development along the colorectal adenoma–carcinoma sequence. Nat Commun, 2015, 6. ArticleID: 6528

[22]

Frey SD, Knorr M, Parrent JL, Simpson RT. Chronic nitrogen enrichment affects the structure and function of the soil microbial community in temperate hardwood and pine forests. For Ecol Manag, 2004, 196(1): 159-171.

[23]

Frey SD, Ollinger S, Nadelhoffer K, Bowden R, Brzostek E, Burton A, Caldwell BA, Crow S, Goodale CL, Grandy AS, Finzi A, Kramer MG, Lajtha K, LeMoine J, Martin M, McDowell WH, Minocha R, Sadowsky JJ, Templer PH, Wickings K. Chronic nitrogen additions suppress decomposition and sequester soil carbon in temperate forests. Biogeochemistry, 2014, 121(2): 305-316.

[24]

Galloway JN, Townsend AR, Erisman JW, Bekunda M, Cai ZC, Freney JR, Martinelli LA, Seitzinger SP, Sutton MA. Transformation of the nitrogen cycle: recent trends, questions, and potential solutions. Science, 2008, 320(5878): 889-892.

[25]

Geng PF, Jin GZ. Fine root morphology and chemical responses to N addition depend on root function and soil depth in a Korean pine plantation in Northeast China. For Ecol Manag, 2022, 520. ArticleID: 120407

[26]

Guan X, Jiang J, Classen AT, Ullah S, Wang GS. Disentangling the contribution of mycorrhizal fungi to soil organic carbon storage. Soil Biol Biochem, 2025, 209. ArticleID: 109900

[27]

He WY, Wei HJ, Liang JW, Xu TY, Chen H, Hu WT, Tang M. How do ectomycorrhizal fungi regulate Eucalyptus biomass allocation through the stoichiometric changes in the plant-soil-microbe system under nitrogen addition?. Ind Crops Prod, 2024, 218. ArticleID: 118876

[28]

He WY, Zhang MM, Jin GZ, Sui X, Zhang T, Song FQ. Effects of nitrogen deposition on nitrogen-mineralizing enzyme activity and soil microbial community structure in a Korean pine plantation. Microb Ecol, 2021, 81(2): 410-424.

[29]

Hendricks JJ, Mitchell RJ, Kuehn KA, Pecot SD. Ectomycorrhizal fungal mycelia turnover in a longleaf pine forest. New Phytol, 2016, 209: 1693-1704.

[30]

Hu JX, Huang CD, Zhou SX, Liu X, Dijkstra FA. Nitrogen addition increases microbial necromass in croplands and bacterial necromass in forests: a global meta-analysis. Soil Biol Biochem, 2022, 165. ArticleID: 108500

[31]

Hu YL, Chen J, Olesen JE, van Groenigen KJ, Hui DF, He XH, Chen GY, Deng Q. Mycorrhizal association controls soil carbon-degrading enzyme activities and soil carbon dynamics under nitrogen addition: a systematic review. Sci Total Environ, 2024, 948. ArticleID: 175008

[32]

Huson DH, Mitra S, Ruscheweyh HJ, Weber N, Schuster SC. Integrative analysis of environmental sequences using MEGAN4. Genome Res, 2011, 21(9): 1552-1560.

[33]

Kallenbach CM, Grandy AS, Frey SD, Diefendorf AF. Microbial physiology and necromass regulate agricultural soil carbon accumulation. Soil Biol Biochem, 2015, 91: 279-290.

[34]

Karlsson FH, Tremaroli V, Nookaew I, Bergström G, Behre CJ, Fagerberg B, Nielsen J, Bäckhed F. Gut metagenome in European women with normal, impaired and diabetic glucose control. Nature, 2013, 498(7452): 99-103.

[35]

Kielak AM, Barreto CC, Kowalchuk GA, van Veen JA, Kuramae EE. The ecology of acidobacteria: moving beyond genes and genomes. Front Microbiol, 2016, 7: 744.

[36]

Lehmann J, Kleber M. The contentious nature of soil organic matter. Nature, 2015, 528(7580): 60-68.

[37]

Lehmann J, Hansel CM, Kaiser C, Kleber M, Maher K, Manzoni S, Nunan N, Reichstein M, Schimel JP, Torn MS, Wieder WR, Kögel-Knabner I. Persistence of soil organic carbon caused by functional complexity. Nat Geosci, 2020, 13(8): 529-534.

[38]

Li JH, Jia HJ, Cai XH, Zhong HZ, Feng Q, Sunagawa S, Arumugam M, Kultima JR, Prifti E, Nielsen T, Juncker AS, Manichanh C, Chen B, Zhang WW, Levenez F, Wang J, Xu X, Xiao L, Liang SS, Zhang DY, Zhang ZX, Chen WN, Zhao HL, Al-Aama JY, Edris S, Yang HM, Wang J, Hansen T, Nielsen HB, Brunak S, Kristiansen K, Guarner F, Pedersen O, Doré J, Ehrlich SD, Bork P, Wang J. An integrated catalog of reference genes in the human gut microbiome. Nat Biotechnol, 2014, 32(8): 834-841.

[39]

Li J, Liu Y, Chen S, Zhou D, Zhang H, Wang J, Li H, Ren Y, Wang C, Song H, Yang X. Soil bacteria drive the pattern of tree regeneration by mediating spatial variation of soil nutrients. J for Res, 2026, 37: 55.

[40]

Liang C, Amelung W, Lehmann J, Kästner M. Quantitative assessment of microbial necromass contribution to soil organic matter. Glob Change Biol, 2019, 25(11): 3578-3590.

[41]

Liang C, Balser TC. Warming and nitrogen deposition lessen microbial residue contribution to soil carbon pool. Nat Commun, 2012, 3. ArticleID: 1222

[42]

Liang C, Schimel JP, Jastrow JD. The importance of anabolism in microbial control over soil carbon storage. Nat Microbiol, 2017, 2. ArticleID: 17105

[43]

Lindahl BD, Tunlid A. Ectomycorrhizal fungi–potential organic matter decomposers, yet not saprotrophs. New Phytol, 2015, 205(4): 1443-1447.

[44]

Liu L, Wen Z, Liu S, Zhang XY, Liu XJ. Decline in atmospheric nitrogen deposition in China between 2010 and 2020. Nat Geosci, 2024, 17(8): 733-736.

[45]

Liu L, Yang JJ, Wang J, Yu Q, Wei CZ, Jiang LC, Huang JH, Zhang YH, Jiang Y, Zhang HY, Han XG. Increase in mineral-associated organic carbon does not offset the decrease in particulate organic carbon under long-term nitrogen enrichment in a steppe ecosystem. Soil Biol Biochem, 2025, 202. ArticleID: 109695

[46]

Lu XF, Hou EQ, Guo JY, Gilliam FS, Li JL, Tang SB, Kuang YW. Nitrogen addition stimulates soil aggregation and enhances carbon storage in terrestrial ecosystems of China: a meta-analysis. Glob Change Biol, 2021, 27(12): 2780-2792.

[47]

Ma T, Zhu SS, Wang ZH, Chen DM, Dai GH, Feng BW, Su XY, Hu HF, Li KH, Han WX, Liang C, Bai YF, Feng XJ. Divergent accumulation of microbial necromass and plant lignin components in grassland soils. Nat Commun, 2018, 9. ArticleID: 3480

[48]

Malik AA, Martiny JBH, Brodie EL, Martiny AC, Treseder KK, Allison SD. Defining trait-based microbial strategies with consequences for soil carbon cycling under climate change. ISME J, 2020, 14(1): 1-9.

[49]

Maynard DS, Crowther TW, Bradford MA. Fungal interactions reduce carbon use efficiency. Ecol Lett, 2017, 20(8): 1034-1042.

[50]

Mielke LA, Klein J, Ekblad A, Finlay RD, Lindahl BD, Clemmensen KE. Fungal guild interactions slow decomposition of boreal forest pine litter and humus. New Phytol, 2025, 247(5): 2367-2380.

[51]

Phillips RP, Brzostek E, Midgley MG. The mycorrhizal-associated nutrient economy: a new framework for predicting carbon–nutrient couplings in temperate forests. New Phytol, 2013, 199(1): 41-51.

[52]

Plett KL, Wojtalewicz D, Anderson IC, Plett JM. Fungal metabolism and free amino acid content may predict nitrogen transfer to the host plant in the ectomycorrhizal relationship between Pisolithus spp. and Eucalyptus grandis. New Phytol, 2024, 242(4): 1589-1602.

[53]

Qiao LK, Wang JF, Wei SS, Ren YL, Lichtfouse E, Han J. The soil microbial carbon pump for carbon sequestration. Environ Chem Lett, 2025, 23(5): 1145-1151.

[54]

Qin JJ, Li RQ, Raes J, Arumugam M, Burgdorf KS, Manichanh C, Nielsen T, Pons N, Levenez F, Yamada T, Mende DR, Li JH, Xu JM, Li SC, Li DF, Cao JJ, Wang B, Liang HQ, Zheng HS, Xie YL, Tap J, Lepage P, Bertalan M, Batto JM, Hansen T, Le Paslier D, Linneberg A, Nielsen HB, Pelletier E, Renault P, Sicheritz-Ponten T, Turner K, Zhu HM, Yu C, Li ST, Jian M, Zhou Y, Li YR, Zhang XQ, Li SG, Qin N, Yang HM, Wang J, Brunak S, Doré J, Guarner F, Kristiansen K, Pedersen O, Parkhill J, Weissenbach J, Bork P, Ehrlich SD, Wang J. A human gut microbial gene catalogue established by metagenomic sequencing. Nature, 2010, 464(7285): 59-65.

[55]

Qin N, Yang FL, Li A, Prifti E, Chen YF, Shao L, Guo J, Le Chatelier E, Yao J, Wu LJ, Zhou JW, Ni SJ, Liu L, Pons N, Batto JM, Kennedy SP, Leonard P, Yuan CH, Ding WC, Chen YT, Hu XJ, Zheng BW, Qian GR, Xu W, Ehrlich SD, Zheng SS, Li LJ. Alterations of the human gut microbiome in liver cirrhosis. Nature, 2014, 513(7516): 59-64.

[56]

Ramirez KS, Craine JM, Fierer N. Consistent effects of nitrogen amendments on soil microbial communities and processes across biomes. Glob Change Biol, 2012, 18(6): 1918-1927.

[57]

Reay DS, Dentener F, Smith P, Grace J, Feely RA. Global nitrogen deposition and carbon sinks. Nat Geosci, 2008, 1(7): 430-437.

[58]

Ren CJ, Zhang XY, Zhang SH, Wang JY, Xu MP, Guo YX, Wang J, Han XH, Zhao FZ, Yang GH, Doughty R. Altered microbial CAZyme families indicated dead biomass decomposition following afforestation. Soil Biol Biochem, 2021, 160. ArticleID: 108362

[59]

Schimel JP, Schaeffer SM. Microbial control over carbon cycling in soil. Front Microbio, 2012, 3: 348.

[60]

Schmidt MWI, Torn MS, Abiven S, Dittmar T, Guggenberger G, Janssens IA, Kleber M, Kögel-Knabner I, Lehmann J, Manning DAC, Nannipieri P, Rasse DP, Weiner S, Trumbore SE. Persistence of soil organic matter as an ecosystem property. Nature, 2011, 478(7367): 49-56.

[61]

Shao SY, Wurzburger N, Sulman B, Hicks Pries C. Ectomycorrhizal effects on decomposition are highly dependent on fungal traits, climate, and litter properties: a model-based assessment. Soil Biol Biochem, 2023, 184. ArticleID: 109073

[62]

Sinsabaugh RL. Phenol oxidase, peroxidase and organic matter dynamics of soil. Soil Biol Biochem, 2010, 42(3): 391-404.

[63]

Six J, Frey SD, Thiet RK, Batten KM. Bacterial and fungal contributions to carbon sequestration in agroecosystems. Soil Sci Soc Am J, 2006, 70(2): 555-569.

[64]

Smith S, Read DJ. Mycorrhizal symbiosis, 2008, third ed, London, Academic Press

[65]

Sokol NW, Sanderman J, Bradford MA. Pathways of mineral-associated soil organic matter formation: integrating the role of plant carbon source, chemistry, and point of entry. Glob Change Biol, 2019, 25(1): 12-24.

[66]

Stevens CJ, Dise NB, Mountford JO, Gowing DJ. Impact of nitrogen deposition on the species richness of grasslands. Science, 2004, 303(5665): 1876-1879.

[67]

Sunagawa S, Coelho LP, Chaffron S, Kultima JR, Labadie K, Salazar G, Djahanschiri B, Zeller G, Mende DR, Alberti A, Cornejo-Castillo FM, Costea PI, Cruaud C, D’ Ovidio F, Engelen S, Ferrera I, Gasol JM, Guidi L, Hildebrand F, Kokoszka F, Lepoivre C, Lima-Mendez G, Poulain J, Poulos BT, Royo-Llonch M, Sarmento H, Vieira-Silva S, Dimier C, Picheral M, Searson S, Kandels-Lewis S, Bowler C, de Vargas C, Gorsky G, Grimsley N, Hingamp P, Iudicone D, Jaillon O, Not F, Ogata H, Pesant S, Speich S, Stemmann L, Sullivan MB, Weissenbach J, Wincker P, Karsenti E, Raes J, Acinas SG, Bork P, Boss E, Bowler C, Follows M, Karp-Boss L, Krzic U, Reynaud EG, Sardet S, Sieracki S, Velayoudon D. Structure and function of the global ocean microbiome. Science, 2015, 348(6237. ArticleID: 1261359

[68]

Terrer C, Jackson RB, Prentice IC, Keenan TF, Kaiser C, Vicca S, Fisher JB, Reich PB, Stocker BD, Hungate BA, Peñuelas J, McCallum I, Soudzilovskaia NA, Cernusak LA, Talhelm AF, Van Sundert K, Piao SL, Newton PCD, Hovenden MJ, Blumenthal DM, Liu YY, Müller C, Winter K, Field CB, Viechtbauer W, Van Lissa CJ, Hoosbeek MR, Watanabe M, Koike T, Leshyk VO, Polley HW, Franklin O. Nitrogen and phosphorus constrain the CO2 fertilization of global plant biomass. Nat Clim Change, 2019, 9(9): 684-689.

[69]

Tian J, Dungait JAJ, Lu XK, Yang YF, Hartley IP, Zhang W, Mo JM, Yu GR, Zhou JZ, Kuzyakov Y. Long-term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil. Glob Change Biol, 2019, 25(10): 3267-3281.

[70]

Ullah MR, Carrillo Y, Dijkstra FA. Drought-induced and seasonal variation in carbon use efficiency is associated with fungi: bacteria ratio and enzyme production in a grassland ecosystem. Soil Biol Biochem, 2021, 155. ArticleID: 108159

[71]

Visscher AM, Vanek S, Meza K, de Goede RGM, Valverde AA, Ccanto R, Olivera E, Scurrah M, Fonte SJ. Eucalyptus and alder field margins differ in their impact on ecosystem services and biodiversity within cropping fields of the Peruvian Andes. Agric Ecosyst Environ, 2020, 303. ArticleID: 107107

[72]

Wang BR, An SS, Liang C, Liu Y, Kuzyakov Y. Microbial necromass as the source of soil organic carbon in global ecosystems. Soil Biol Biochem, 2021, 162. ArticleID: 108422

[73]

Wang Z, Yang L, Wang J, Zhao X, Zhang C, von Gadow K. Soil fertility and forest structure influence tree carbon stock depending on mycorrhizal types in a temperate forest. J for Res, 2025, 36: 115.

[74]

Wieder WR, Allison SD, Davidson EA, Georgiou K, Hararuk O, He YJ, Hopkins F, Luo YQ, Smith MJ, Sulman B, Todd-Brown K, Wang YP, Xia JY, Xu XF. Explicitly representing soil microbial processes in Earth system models. Glob Biogeochem Cycles, 2015, 29(10): 1782-1800.

[75]

Willard SJ, Vanguelova E, Waring B. Nitrogen deposition is linked to changes in mineral-associated organic carbon in forest soils in the United Kingdom. Plant Soil, 2026, 519(2): 2051-2069.

[76]

Wu H, Yang JJ, Fu W, Rillig MC, Cao ZJ, Zhao AH, Hao ZP, Zhang X, Chen BD, Han XG. Identifying thresholds of nitrogen enrichment for substantial shifts in arbuscular mycorrhizal fungal community metrics in a temperate grassland of Northern China. New Phytol, 2023, 237(1): 279-294.

[77]

Wu JJ, Zhang H, Pan YT, Cheng XL, Zhang KR, Liu GH. Particulate organic carbon is more sensitive to nitrogen addition than mineral-associated organic carbon: a meta-analysis. Soil Tillage Res, 2023, 232. ArticleID: 105770

[78]

Wu R, Zhang H, Zhang M, Jin G, Song F. Saprotrophic fungal community responses to nitrogen additions in a Korean pine plantation: insights from using the mycoindicator. J for Res, 2025, 36: 27.

[79]

Xiong MY, Jiang W, Zou SZ, Kang D, Yan XC. Microbial carbohydrate-active enzymes influence soil carbon by regulating the of plant- and fungal-derived biomass decomposition in plateau peat wetlands under differing water conditions. Front Microbiol, 2023, 14. ArticleID: 1266016

[80]

Xu FD, Li C, Chen YX, Wu JC, Bai HD, Fan SG, Yang YC, Zhang YP, Li SF, Su JR. Soil microbial community structure and soil fertility jointly regulate soil microbial residue carbon during the conversion from subtropical primary forest to plantations. Geoderma, 2024, 441. ArticleID: 116767

[81]

Xu JR, Zhu WL, Yu SH, Dun XJ, Hu DM, Liao DX, Hao M, Lian FJ, Zhang ZX, Gao P. Effects of thinning on soil microbial community and carbon fractions and their relationships in coastal protected forests. Environ Earth Sci, 2025, 84(8): 207.

[82]

Yang LY, Canarini A, Zhang WS, Lang M, Chen YX, Cui ZL, Kuzyakov Y, Richter A, Chen XP, Zhang FS, Tian J. Microbial life-history strategies mediate microbial carbon pump efficacy in response to N management depending on stoichiometry of microbial demand. Glob Change Biol, 2024, 30(5. ArticleID: e17311

[83]

Yu GR, Jia YL, He NP, Zhu JX, Chen Z, Wang QF, Piao SL, Liu XJ, He HL, Guo XB, Wen Z, Li P, Ding GA, Goulding K. Stabilization of atmospheric nitrogen deposition in China over the past decade. Nat Geosci, 2019, 12(6): 424-429.

[84]

Yu YL, Zhu NX, Ren Y, Dong MH, Sun GF, Virk AL, Li FM, Yang HS, Kan ZR. Effects of crop rotation on plant- and microbial-derived carbon within particulate and mineral fractions in paddy soils. Agric Ecosyst Environ, 2025, 380. ArticleID: 109398

[85]

Yuan X, Qin WK, Xu H, Zhang ZH, Zhou HK, Zhu B. Sensitivity of soil carbon dynamics to nitrogen and phosphorus enrichment in an Alpine meadow. Soil Biol Biochem, 2020, 150. ArticleID: 107984

[86]

Zhang MM, Lu Y, Jin GZ, Zhu B. Bacterial communities in litter are more sensitive to high nitrogen addition than fungal communities in a Korean pine (Pinus koraiensis) plantation. J For Res, 2026, 37(1): 23.

[87]

Zhang XY, Jia J, Chen LT, Chu HY, He JS, Zhang YJ, Feng XJ. Aridity and NPP constrain contribution of microbial necromass to soil organic carbon in the Qinghai-Tibet Alpine grasslands. Soil Biol Biochem, 2021, 156. ArticleID: 108213

[88]

Zhu XF, Jackson RD, DeLucia EH, Tiedje JM, Liang C. The soil microbial carbon pump: from conceptual insights to empirical assessments. Glob Change Biol, 2020, 26(11): 6032-6039.

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