doi:10.48130/fra-0026-0015

[1]

Lehmann J, Kleber M. 2015. The contentious nature of soil organic matter. Nature 528:60−68

doi: 10.1038/nature16069

[2]

Stockmann U, Adams MA, Crawford AE, Field DJ, Henakaarchchi M, et al. 2013. The knowns, known unknowns and unknowns of sequestration of soil organic carbon. Agriculture, Ecosystems and Environment 164:80−99

doi: 10.1016/j.agee.2012.10.001

[3]

Soudzilovskaia NA, van der Heijden MGA, Cornelissen JHC, Makarov MI, Onipchenko VG, et al. 2015. Quantitative assessment of the differential impacts of arbuscular and ectomycorrhiza on soil carbon cycling. New Phytologist 208:280−293

doi: 10.1111/nph.13447

[4]

Frey SD. 2019. Mycorrhizal fungi as mediators of soil organic matter dynamics. Annual Review of Ecology, Evolution, and Systematics 50:237−259

doi: 10.1146/annurev-ecolsys-110617-062331

[5]

Högberg MN, Briones MJI, Keel SG, Metcalfe DB, Campbell C, et al. 2010. Quantification of effects of season and nitrogen supply on tree below-ground carbon transfer to ectomycorrhizal fungi and other soil organisms in a boreal pine forest. New Phytologist 187:485−493

doi: 10.1111/j.1469-8137.2010.03274.x

[6]

Soudzilovskaia NA, van Bodegom PM, Terrer C, Zelfde MV, McCallum I, et al. 2019. Global mycorrhizal plant distribution linked to terrestrial carbon stocks. Nature Communications 10:5077

doi: 10.1038/s41467-019-13019-2

[7]

Taylor MK, Lankau RA, Wurzburger N. 2016. Mycorrhizal associations of trees have different indirect effects on organic matter decomposition. Journal of Ecology 104:1576−1584

doi: 10.1111/1365-2745.12629

[8]

Fernandez CW, Kennedy PG. 2015. Moving beyond the black-box: fungal traits, community structure, and carbon sequestration in forest soils. New Phytologist 205:1378−1380

doi: 10.1111/nph.13289

[9]

Hendricks JJ, Mitchell RJ, Kuehn KA, Pecot SD, Sims SE. 2006. Measuring external mycelia production of ectomycorrhizal fungi in the field: the soil matrix matters. New Phytologist 171:179−186

doi: 10.1111/j.1469-8137.2006.01742.x

[10]

Nilsson LO, Giesler R, Bååth E, Wallander H. 2005. Growth and biomass of mycorrhizal mycelia in coniferous forests along short natural nutrient gradients. New Phytologist 165:613−622

doi: 10.1111/j.1469-8137.2004.01223.x

[11]

Nilsson LO, Wallander H. 2003. Production of external mycelium by ectomycorrhizal fungi in a norway spruce forest was reduced in response to nitrogen fertilization. New Phytologist 158:409−416

doi: 10.1046/j.1469-8137.2003.00728.x

[12]

Wurzburger N, Jack Brookshire ENJ. 2017. Experimental evidence that mycorrhizal nitrogen strategies affect soil carbon. Ecology 98:1491−1497

doi: 10.1002/ecy.1827

[13]

Yin L, Dijkstra FA, Phillips RP, Zhu B, Wang P, et al. 2021. Arbuscular mycorrhizal trees cause a higher carbon to nitrogen ratio of soil organic matter decomposition via rhizosphere priming than ectomycorrhizal trees. Soil Biology and Biochemistry 157:108246

doi: 10.1016/j.soilbio.2021.108246

[14]

Wu S, Fu W, Rillig MC, Chen B, Zhu Y, et al. 2024. Soil organic matter dynamics mediated by arbuscular mycorrhizal fungi–an updated conceptual framework. New Phytologist 242:1417−1425

doi: 10.1111/nph.19178

[15]

Klink S, Keller AB, Wild AJ, Baumert VL, Gube M, et al. 2022. Stable isotopes reveal that fungal residues contribute more to mineral associated organic matter pools than plant residues. Soil Biology and Biochemistry 168:108634

doi: 10.1016/j.soilbio.2022.108634

[16]

Smith SE, Read DJ. 2008. Mycorrhizal Symbiosis. 3rd edition. Cambridge, MA: Elsevier, Academic Press. pp. 1−787 doi: 10.1016/B978-0-12-370526-6.X5001-6

[17]

Ward EB, Duguid MC, Kuebbing SE, Lendemer JC, et al. 2021. The functional role of ericoid mycorrhizal plants and fungi on carbon and nitrogen dynamics in forests. New Physiologist 235:1701−1718

doi: 10.1111/nph.18307

[18]

Read DJ, Perez-Moreno J. 2003. Mycorrhizas and nutrient cycling in ecosystems – a journey towards relevance? New Phytologist 157:475−492

doi: 10.1046/j.1469-8137.2003.00704.x

[19]

Conti G, Urcelay C, Gundel PE, Piñeiro G. 2025. The potential of arbuscular mycorrhizal fungi to improve soil organic carbon in agricultural ecosystems: a meta-analytical approach. Functional Ecology 39:1016−1030

doi: 10.1111/1365-2435.14753

[20]

Nicolás C, Martin-Bertelsen T, Floudas D, Bentzer J, Smits M, et al. 2019. The soil organic matter decomposition mechanisms in ectomycorrhizal fungi are tuned for liberating soil organic nitrogen. The ISME Journal 13:977−988

doi: 10.1038/s41396-018-0331-6

[21]

Clemmensen KE, Bahr A, Ovaskainen O, Dahlberg A, Ekblad A, et al. 2013. Roots and associated fungi drive long-term carbon sequestration in boreal forest. Science 339:1615−1618

doi: 10.1126/science.1231923

[22]

Clemmensen KE, Finlay RD, Dahlberg A, Stenlid J, Wardle DA, et al. 2015. Carbon sequestration is related to mycorrhizal fungal community shifts during long-term succession in boreal forests. New Phytologist 205:1525−1536

doi: 10.1111/nph.13208

[23]

Tedersoo L, Bahram M. 2019. Mycorrhizal types differ in ecophysiology and alter plant nutrition and soil processes. Biological Reviews 94:1857−1880

doi: 10.1111/brv.12538

[24]

See CR, Keller AB, Hobbie SE, Kennedy PG, Weber PK, et al. 2022. Hyphae move matter and microbes to mineral microsites: integrating the hyphosphere into conceptual models of soil organic matter stabilization. Global Change Biology 28:2527−2540

doi: 10.1111/gcb.16073

[25]

Terrer C, Vicca S, Hungate BA, Phillips RP, Prentice IC. 2016. Mycorrhizal association as a primary control of the CO₂ fertilization effect. Science 353:72−74

doi: 10.1126/science.aaf46

[26]

Treseder KK, Allen MF. 2000. Mycorrhizal fungi have a potential role in soil carbon storage under elevated CO₂ and nitrogen deposition. New Phytologist 147:189−200

doi: 10.1046/j.1469-8137.2000.00690.x

[27]

Näsholm T, Högberg P, Franklin O, Metcalfe D, Keel SG, et al. 2013. Are ectomycorrhizal fungi alleviating or aggravating nitrogen limitation of tree growth in boreal forests? New Phytologist 198:214−221

doi: 10.1111/nph.12139

[28]

Davidson EA, Janssens IA. 2006. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature 440:165−173

doi: 10.1038/nature04514

[29]

Serra-Maluquer X, Mencuccini M, Martínez-Vilalta J. 2018. Changes in tree resistance, recovery and resilience across three successive extreme droughts in the northeast Iberian Peninsula. Oecologia 187:343−354

doi: 10.1007/s00442-018-4118-2

[30]

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

doi: 10.1038/nmicrobiol.2017.105

[31]

Liang C, Kästner M, Joergensen RG. 2020. Microbial necromass on the rise: the growing focus on its role in soil organic matter development. Soil Biology and Biochemistry 150:108000

doi: 10.1016/j.soilbio.2020.108000

[32]

Finzi AC, Abramoff RZ, Spiller KS, Brzostek ER, Darby BA, et al. 2015. Rhizosphere processes are quantitatively important components of terrestrial carbon and nutrient cycles. Global Change Biology 21:2082−2094

doi: 10.1111/gcb.12816

[33]

Meier IC, Prichard SG, Brzostek ER, McCormack ML, Phillips RP. 2015. The rhizosphere and hyphosphere differ in their impacts on carbon and nitrogen cycling in forests exposed to elevated CO₂. New Phytologist 205:1164−1174

doi: 10.1111/nph.13122

[34]

Villarino SH, Pinto P, Jackson RB, Piñeiro G. 2021. Plant rhizodeposition: a key factor for soil organic matter formation in stable fractions. Science Advances 7:eabd3176

doi: 10.1126/sciadv.abd3176

[35]

Fernandez CW, Kennedy PG. 2016. Revisiting the 'Gadgil effect': do interguild fungal interactions control carbon cycling in forest soils? New Phytologist 209:1382−1394

doi: 10.1111/nph.13648

[36]

Hawkins HJ, Cargill RIM, Van Nuland ME, Hagen SC, Field KJ, et al. 2023. Mycorrhizal mycelium as a global carbon pool. Current Biology 33:R560−R573

doi: 10.1016/j.cub.2023.02.027

[37]

Van't Padje A, Werner GDA, Kiers ET. 2021. Mycorrhizal fungi control phosphorus value in trade symbiosis with host roots when exposed to abrupt 'crashes' and 'booms' of resource availability. New Phytologist 229:2933−2944

doi: 10.1111/nph.17055

[38]

Cairney JWG. 2012. Extramatrical mycelia of ectomycorrhizal fungi as moderators of carbon dynamics in forest soil. Soil Biology and Biochemistry 47:198−208

doi: 10.1016/j.soilbio.2011.12.029

[39]

Angst G, Mueller KE, Nierop KGJ, Simpson MJ. 2021. Plant-or microbial- derived? a review on the molecular composition of stabilized soil organic matter. Soil Biology and Biochemistry 156:108189

doi: 10.1016/j.soilbio.2021.108189

[40]

Adamczyk B, Sietiö OM, Straková P, Prommer J, Wild B, et al. 2019. Plant roots increase both decomposition and stable organic matter formation in boreal forest soil. Nature Communications 10:3982

doi: 10.1038/s41467-019-11993-1

[41]

Dijkstra FA, Zhu B, Cheng W. 2021. Root effects on soil organic carbon: a double-edged sword. New Phytologist 230:60−65

doi: 10.1111/nph.17082

[42]

Lindahl BD, Tunlid A. 2015. Ectomycorrhizal fungi − potential organic matter decomposers, yet not saprotrophs. New Phytologist 205:1443−1447

doi: 10.1111/nph.13201

[43]

Dudley MP, Freeman M, Wenger S, Jackson CR, Pringle CM. 2020. Rethinking foundation species in a changing world: the case for Rhododendron maximum as an emerging foundation species in shifting ecosystems of the southern Appalachians. Forest Ecology and Management 472:118240

doi: 10.1016/j.foreco.2020.118240

[44]

Averill C, Hawkes CV. 2016. Ectomycorrhizal fungi slow soil carbon cycling. Ecology Letters 19:937−947

doi: 10.1111/ele.12631

[45]

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

doi: 10.1038/nature12901

[46]

Rillig MC, Mummey DL. 2006. Mycorrhizas and soil structure. New Phytologist 171:41−53

doi: 10.1111/j.1469-8137.2006.01750.x

[47]

Beidler KV, Phillips RP, Andrews E, Maillard F, Mushinski RM, et al. 2020. Substrate quality drives fungal necromass decay and decomposer community structure under contrasting vegetation types. Journal of Ecology 108:1845−1859

doi: 10.1111/1365-2745.13385

[48]

Fei S, Kivlin SN, Domke GM, Jo I, LaRue EA, et al. 2022. Coupling of plant and mycorrhizal fungal diversity: its occurrence, relevance, and possible implications under global change. New Phytologist 236:1960−1966

doi: 10.1111/nph.17954

[49]

Ekblad A, Wallander H, Godbold DL, Cruz C, Johnson D, et al. 2013. The production and turnover of extramatrical mycelium of ectomycorrhizal fungi in forest soils: role in carbon cycling. Plant and Soil 366:1−27

doi: 10.1007/s11104-013-1630-3

[50]

Netherway T, Bahram M. 2024. Melanized root-associated fungi: key players in plant–soil systems. Trends in Microbiology 32:1190−1199

doi: 10.1016/j.tim.2024.06.006

[51]

Coelho RRR, Sacramento DR, Linhares LF. 1997. Amino sugars in fungal melanins and soil humic acids. European Journal of Soil Science 48: 425–429

[52]

Zhou J, Zang H, Loeppmann S, Gube M, Kuzyakov Y, et al. 2020. Arbuscular mycorrhiza enhances rhizodeposition and reduces the rhizosphere priming effect on the decomposition of soil organic matter. Soil Biology and Biochemistry 140:107641

doi: 10.1016/j.soilbio.2019.107641

[53]

Jeewani PH, Luo Y, Yu G, Fu Y, He X, et al. 2021. Arbuscular mycorrhizal fungi and goethite promote carbon sequestration via hyphal-aggregate mineral interactions. Soil Biology and Biochemistry 162:108417

doi: 10.1016/j.soilbio.2021.108417

[54]

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

doi: 10.1111/nph.12221

[55]

Lang AK, Jevon FV, Vietorisz CR, Ayres MP, Hatala Matthes J. 2021. Fine roots and mycorrhizal fungi accelerate leaf litter decomposition in a northern hardwood forest regardless of dominant tree mycorrhizal associations. New Phytologist 230:316−326

doi: 10.1111/nph.17155

[56]

Zhu K, McCormack ML, Lankau RA, Egan JF, Wurzburger N. 2018. Association of ectomycorrhizal trees with high carbon-to nitrogen ratio soils across temperate forests is driven by smaller nitrogen not larger carbon stocks. Journal of Ecology 106:524−535

doi: 10.1111/1365-2745.12918

[57]

Fitch AA, Lang AK, Whalen ED, Geyer K, Hicks Pries C. 2020. Fungal community, not substrate quality, drives soil microbial function in northeastern U.S. temperate forests. Frontiers in Forests and Global Change 3:1−13

doi: 10.3389/ffgc.2020.569945

[58]

Lavallee JM, Soong JL, Cotrufo MF. 2020. Conceptualizing soil organic matter into particulate and mineral-associated forms to address global change in the 21^st century. Global Change Biology 26:261−273

doi: 10.1111/gcb.14859

[59]

Rillig MC, Aguilar-Trigueros CA, Bergmann J, Verbruggen E, Veresoglou SD, et al. 2015. Plant root and mycorrhizal fungal traits for understanding soil aggregation. New Phytologist 205:1385−1388

doi: 10.1111/nph.13045

[60]

Burford EP, Kierans M, Gadd GM. 2003. Geomycology: fungi in mineral substrata. Mycologist 17:98−107

doi: 10.1017/S0269915X03003112

[61]

Hemingway JD, Rothman DH, Grant KE, Rosengard SZ, Eglinton TI, et al. 2019. Mineral protection regulates long-term global preservation of natural organic carbon. Nature 570:228−231

doi: 10.1038/s41586-019-1280-6

[62]

Qiao Y, Miao S, Li N, Han X, Zhang B. 2014. Spatial distribution of rhizodeposit carbon of maize (Zea mays L. ) in soil aggregates assessed by multiple pulse ¹³C labeling in the field. Plant and Soil 375:317−329

doi: 10.1007/s11104-013-1932-5

[63]

Zhu X, Jackson RD, De Lucia EH, Tiedje JM, Liang C. 2020. The soil microbial carbon pump: from conceptual insights to empirical assessments. Global Change Biology 26:6032−6039

doi: 10.1111/gcb.15319

[64]

Perotto S, Daghino S, Martino E. 2018. Ericoid mycorrhizal fungi and their genomes: another side to the mycorrhizal symbiosis? New Phytologist 220:1141−1147

doi: 10.1111/nph.15218

[65]

Wurzburger N, Hendrick RL. 2009. Plant litter chemistry and mycorrhizal roots promote a nitrogen feedback in a temperate forest. Journal of Ecology 97:528−536

doi: 10.1111/j.1365-2745.2009.01487.x

[66]

Brundrett MC, Tedersoo L. 2018. Evolutionary history of mycorrhizal symbioses and global host plant diversity. New Phytologist 220:1108−1115

doi: 10.1111/nph.14976

[67]

Karlen DL, Cambardella CA. 2020. Conservation strategies for improving soil quality and organic matter storage. In Structure and Organic Matter Storage in Agricultural Soils. eds. Carter MR, Stewart BA. Boca Raton: CRC Press. pp. 395−420 doi: 10.1201/9781003075561-15

[68]

Chowdhury S, Lange M, Malik AA, Goodall T, Huang J, et al. 2022. Plants with arbuscular mycorrhizal fungi efficiently acquire nitrogen from substrate additions by shaping the decomposer community composition and their net plant carbon demand. Plant and Soil 475:473−490

doi: 10.1007/s11104-022-05380-x

[69]

Andrino A, Boy J, Mikutta R, Sauheitl L, Guggenberger G. 2019. Carbon investment required for the mobilization of inorganic and organic phosphorus bound to goethite by an arbuscular mycorrhiza (Solanum lycopersicum × Rhizophagus irregularis). Frontiers in Environmental Science 7:26

doi: 10.3389/fenvs.2019.00026

[70]

Daynes CN, Field DJ, Saleeba JA, Cole MA, McGee PA. 2013. Development and stabilization of soil structure via interactions between organic matter, arbuscular mycorrhizal fungi, and plant roots. Soil Biology and Biochemistry 57:683−694

doi: 10.1016/j.soilbio.2012.09.020

[71]

Cheng L, Booker FL, Tu C, Burkey KO, Zhou L, et al. 2012. Arbuscular mycorrhizal fungi increase organic carbon decomposition under elevated CO₂. Science 337:1084−1087

doi: 10.1126/science.1224304

[72]

Tao J, Liu X. 2024. Does arbuscular mycorrhizal fungi inoculation influence soil carbon sequestration? Biology and Fertility of Soils 60:213−225

doi: 10.1007/s00374-024-01793-1

[73]

Six J, Bossuyt H, Degryze S, Denef K. 2004. A history of research on the link between (micro) aggregates, soil biota, and soil organic matter dynamics. Soil and Tillage Research 79:7−31

doi: 10.1016/j.still.2004.03.008

[74]

Jeffries P, Gianinazzi S, Perotto S, Turnau K, Barea JM. 2003. The contribution of arbuscular mycorrhizal fungi in sustainable maintenance of plant health and soil fertility. Biology and Fertility of Soils 37:1−16

doi: 10.1007/s00374-002-0546-5

[75]

Marro N, Cofré N, Grilli G, Alvarez C, Labuckas D, et al. 2020. Soybean yield, protein content and oil quality in response to interaction of arbuscular mycorrhizal fungi and native microbial populations from mono- and rotation-cropped soils. Applied Soil Ecology 152:103575

doi: 10.1016/j.apsoil.2020.103575

[76]

Langley JA, Hungate BA. 2003. Mycorrhizal controls on belowground litter quality. Ecology 84:2302−2312

doi: 10.1890/02-0282

[77]

Cotrufo MF, Wallenstein MD, Boot CM, Denef K, Paul E. 2013. 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? Global Change Biology 19:988−995

doi: 10.1111/gcb.12113

[78]

Treseder KK, Lennon JT. 2015. Fungal traits that drive ecosystem dynamics on land. Microbiology and Molecular Biology Reviews 79:243−262

doi: 10.1128/mmbr.00001-15

[79]

Talbot JM, Bruns TD, Smith DP, Branco S, Glassman SI, et al. 2013. Independent roles of ectomycorrhizal and saprotrophic communities in soil organic matter decomposition. Soil Biology and Biochemistry 57:282−291

doi: 10.1016/j.soilbio.2012.10.004

[80]

Clemmensen KE, Durling MB, Michelsen A, Hallin S, Finlay RD, et al. 2021. A tipping point in carbon storage when forest expands into tundra is related to mycorrhizal recycling of nitrogen. Ecology Letters 24:1193−1204

doi: 10.1111/ele.13735

[81]

See CR, Luke McCormack M, Hobbie SE, Flores-Moreno H, Silver WL, et al. 2019. Global patterns in fine root decomposition: climate, chemistry, mycorrhizal association and woodiness. Ecology Letters 22:946−953

doi: 10.1111/ele.13248

[82]

Adamczyk B, Sietiö OM, Biasi C, Heinonsalo J. 2019. Interaction between tannins and fungal necromass stabilizes fungal residues in boreal forest soils. New Phytologist 223:16−21

doi: 10.1111/nph.15729

[83]

Read DJ. 1991. Mycorrhizas in ecosystems. Experientia 47:376−391

doi: 10.1007/BF01972080

[84]

Six J, Frey SD, Thiet RK, Batten KM. 2006. Bacterial and fungal contributions to carbon sequestration in agroecosystems. Soil Science Society of America Journal 70:555−569

doi: 10.2136/sssaj2004.0347

[85]

Terrer C, Phillips RP, Hungate BA, Rosende J, Pett-Ridge J, et al. 2021. A trade-off between plant and soil carbon storage under elevated CO₂. Nature 591:599−603

doi: 10.1038/s41586-021-03306-8

[86]

Xu Z, Hu R, Xiong P, Wan C, Cao G, et al. 2010. Initial soil responses to experimental warming in two contrasting forest ecosystems, Eastern Tibetan Plateau, China: Nutrient availabilities, microbial properties and enzyme activities. Applied Soil Ecology 46:291−299

doi: 10.1016/j.apsoil.2010.07.005

[87]

Qiu Y, Zhao Y, Wang B, Xu X, He T, et al. 2026. Root traits and mycorrhizal fungi mediate reactive N and warming impacts on soil organic carbon. Nature Communications 17:3184

doi: 10.1038/s41467-026-69301-7

[88]

Fernandez CW, Heckman K, Kolka R, Kennedy PG. 2019. Melanin mitigates the accelerated decay of mycorrhizal necromass with peatland warming. Ecology Letters 22:498−505

doi: 10.1111/ele.13209

[89]

Odriozola I, Větrovský T, Barbi F, Machac A, Dobbler PT, et al. 2026. Global distribution and biogeography of ericoid mycorrhizal fungi. New Phytologist 251:2135−2151

doi: 10.1111/nph.71334

[90]

Cao JL, Xie L, Zheng Y, Yang Y. 2020. Drought intensify the effects of warming on root-colonizing arbuscular mycorrhizal fungal community in subtropical Chinese fir plantation. Forest Ecology and Management 464:118078

doi: 10.1016/j.foreco.2020.118078

[91]

Hagenbo A, Piñuela Y, Castaño C, Martínez de Aragón J, de-Miguel S, et al. 2021. Production and turnover of mycorrhizal soil mycelium relate to variation in drought conditions in Mediterranean Pinus pinaster, Pinus sylvestris and Quercus ilex forests. New Phytologist 230:1609−1622

doi: 10.1111/nph.17012

[92]

Deng L, Peng C, Kim DG, Liu Y, Hai X, et al. 2021. Drought effects on soil carbon and nitrogen dynamics in global natural ecosystems. Earth-Science Reviews 214:103501

doi: 10.1016/j.earscirev.2020.103501

[93]

Bradford MA, Reynolds JF. 2006. Scaling terrestrial biogeochemical processes: contrasting intact and model experimental systems. In Scaling and uncertainty analysis in ecology: methods and applications, eds. Wu J, Jones B, Li H, Loucks OL. Dordrecht: Springer. pp. 107–128 doi: 10.1007/1-4020-4663-4_6